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ISSN 0073-6767 

Proceedings of the 

Indiana Academy 
of Science 

Volume 106 

Number 1-2 

The PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE is a quarterly journal dedicated to pro- 
moting scientific research and the diffusion of scientific information; to encouraging communication and 
cooperation among scientists; and to improving education in the sciences. 


Gary E. Dolph 
Indiana University Kokomo 

2300 S. Washington St. 

Kokomo, Indiana 46904-9003 




Hans O. Andersen 
Indiana University 

Rita Ban- 
Purdue University 

Ernest E. Campaigne 
Indiana University 

William R. Clark 
Ball State University 

Donald R. Cochran 
Ball State University 

Robert F. Dale 
Purdue University 

Kara W. Eberly 
St. Mary's College 

Uwe J. Hansen 
Indiana State University 

Darly R. Karns 
Hanover College 

N. Gary Lane 
Indiana University 

Paul C. MacMillan 
Hanover College 

James D. Haddock 
Rebecca Dolan 
Susan M. Johnson 
Edward L. Frazier 
James W. Berry 
Gary E. Dolph 
James Gammon 
Nelson R. Shaffer 

Wilton N. Melhorn 
Purdue University 

Paul Rothrock 
Taylor University 

Alfred R. Shmidt 

Rose Hulman Institute of Technology 

Thomas P. Simon 

United States Environmental Protection Agency 

Paul M. Stewart 

Indiana University-Purdue University Fort Wayne 

Michael Tansey 
Indiana University 

Robert D. Waltz 

Indiana Department of Natural Resources 

J. Dan Webster 
Hanover College 

Harmon Weeks 
Purdue University 

John O. Whitaker, Jr. 
Indiana State University 


Indiana University-Purdue Univ. Fort Wayne 

Butler University 

Ball State University 

5007 W. 14th Street, Speedway 

Butler University 

Indiana University Kokomo 

DePauw University 

Indiana Geological Survey 





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Founded 29 December 1885 


Volume 106 Number 1-2 

Published at Bloomington, Indiana 
April 15, 1999 

Copyright © 1999 by the Indiana Academy of Science 


Volume 106, No. 1-2(1997) 









S.E. Brown and G.R. Parker. IMPACT OF WHITE-TAILED 

T.P Simon, R.N. Jankowski, and C. Morris. PHYSICAL AND 

1980-1995 67 

J.O. Whitaker, Jr., R. McKenzie, M. Rakow, B. Leibacher, and 


R.A. Cloyd, C.R. Edwards, and L.W. Bledsoe. TRICHOME 



J. Pichtel, A. Covey, and K. Lukscay. REMOVAL OF LEAD 




H.G. Day. HERMAN T. BRISCOE (1893-1960): A SUPERIOR 


E.M. Ossom, C.U. Ethothi, and C.L. Rhykerd. INFLUENCE OF 



Proceedings of the Indiana Academy of Science 1 

(1997) Volume 106 p. 1-23 





Raelene M. Crandall and Rebecca W. Dolan 

Department of Biological Sciences 

Butler University 

Indianapolis, Indiana 46208 

ABSTRACT: The protection of plant resources in urban areas is a growing 
conservation concern. Inventory activities that document species presence and 
stewardship plans that protect and enhance these areas are needed. The results 
of a botanical inventory of the Crooked Creek Community Juan Solomon Park 
in Indianapolis, Indiana, are reported in this paper. The 46-acre park con- 
tains three distinct habitats, supporting a wide variety of plants. One hun- 
dred seventy-nine vascular plant species from 64 families were identified, 
including 53 (29.6%) non-native species that are naturalizing within the park. 
Despite its high percentage of alien species and urban setting, the park is an 
important natural area. The flora's coefficient of conservatism (sensu Swink 
and Wilhelm, 1994) was 54.1. Several exotic, invasive species (most 
notably garlic mustard, amur bush honeysuckle, and wintercreeper) pose poten- 
tial future threats to the park's natural flora, and management efforts should 
be focused on their removal. 

KEYWORDS: Coefficient of conservatism, floristics, Indiana, invasive exotics. 


Floristic investigations that document existing plant resources provide essen- 
tial information for the sound stewardship of natural areas. Management rec- 
ommendations based on inventory studies can help maintain natural areas through 
the protection of high-quality habitats and the control of invasive, exotic species. 
A botanical exploration of Crooked Creek Community Juan Solomon Park (a 
park run by the Indianapolis Parks Department) was carried out during the flow- 
ering season of 1996. The vascular plant species in the park were identified, and 
voucher specimens were collected. The major plant communities and habitats 
were also identified. The value of the park as a natural area was assessed using 
Swink and Wilhelm's (1994) coefficient of conservatism. Control recommen- 
dations were developed for potentially invasive species located in the park. 

Crooked Creek Community Juan Solomon Park is located at the northwest 
corner of Grandview Drive and Fox Hill Road along Crooked Creek in Wash- 
ington Township, Marion County, Indianapolis, Indiana (Lat. 39° 51 ' N, Long. 
86° 11' W; Sec. 3, T16N, R3E, Indianapolis West Quadrangle). The park con- 
sists of 46 acres. Twenty-four acres were purchased by the city in 1975, and the 
land provides recreational and educational opportunities for many of the local 
residents. An additional 22 acres were purchased by the Crooked Creek Com- 

Botany: Crandall and Dolan 

Vol. 106 (1997) 

Figure 1. The major plant communities and habitats studied at the Crooked Creek 
Community Juan Solomon Park. 

munity Council and donated to the city in September 1995. This area consists 
primarily of mesic upland woods. Crooked Creek, a fairly pristine creek, bisects 
the park into two roughly equal halves. Large sycamores and other hardwoods 
line the creek. 

Although located in the most populous county in the State, Crooked Creek 
Community Juan Soloman Park is situated in a landscape of housing subdivi- 
sions and recently abandoned fields. The soils are mostly Genesee silt loams: 
deep, nearly level, well-drained soils formed in loamy alluvium (Sturm and 
Gilbert, 1978). Along the creek, small sand bars and sand spots were mapped 
within the soil. Indianapolis is located in the Tipton Till Plain Natural Region 
(sensu Homoya, et al., 1985), a region of primarily undissected plain formerly 
covered by beech-maple-oak forest. 


To conduct the floristic survey, the park was divided into 5 sections 
according to geographical landscape (Figure 1). Each section was repeatedly vis- 
ited to identify and collect as many different plants as possible. Specimens were 
collected over five months from April, 1996, to September, 1996, with the great- 
est emphasis during the months of June and July, the peak flowering season. The 
park was visited approximately 3 times a week during June and July and at least 
every two weeks during the remaining months. 

Vol. 106 (1997) Indiana Academy of Science 3 

Plant species were collected only if they were not considered rare, threat- 
ened, or endangered and if the sampling site had more than 20 individuals. Vouch- 
er specimens were deposited in the Friesner Herbarium (BUT) of Butler University, 
Indianapolis, Indiana. Photographic documentation is available for the majori- 
ty of the species that could not be collected due to small sample size, inacess- 
ability of the leaves from large trees, or the adverse effects of the plant on the 
collector (stinging nettle or poison ivy). Plants were identified using published 
reference manuals. The nomenclature follows Gleason and Cronquist (1991). 
The Indiana Department of Natural Resource's publication (1993) on Indiana's 
rare plants and animals was used to identify rare, threatened, or endangered 
plants. In addition, management recommendations for the control and identifi- 
cation of invasive exotics were developed. 

The coefficient of conservatism (Swink and Wilhelm, 1994) was calculat- 
ed for the flora to determine the park's quality as a natural area. Swink and 
Wilhelm (1994) determined C values by examining the fidelity of species to high 
quality habitats. For this calculation, native species were assigned quality 
index values (C), ranging from for species that are not habitat specialists to 10 
for species that are indicators of high-quality plant communities. 

The coefficient of conservatism (I) was calculated using the equation 

i = cVn 

where C represents the average quality index value of all the native species 
present, and N is the number of native species. 

Based on site surveys in the Chicago area, Swink and Wilhelm (1994) 
rated sites with I values of less than 35 as not significant from a natural areas 
perspective, sites with I values from 35 to 50 as significant, and sites with I 
values greater than 50 as of paramount importance for conservation. The coef- 
ficient of conservatism is not based on the abundance or frequency of the native 
species, because these values can vary with the seasons or between years. In 
addition, exotic plant species are not used in the calculation. Swink and Wilhelm 
(1994) believe that a low coefficient of conservatism indicates the negative impact 
of non-native species; only the native species present are used to determine of 
the quality of the flora using their scheme. However, this system was developed 
for the Chicago area. Although all but two native species found in this study were 
given quality index values by Swink and Wilhelm, the habitat fidelity charac- 
teristics of the plants may be different in the park, which is located ca. 325 km 
south of Chicago. Efforts have been made to develop this methodology for other 
geographical regions, but no system is currently available for central Indiana 
(Herman, 1997; Ladd, in prep.). 


Based on repeated visits to the five survey sections of the park, three dis- 
tinct habitats were identied: mesic upland woods, lowland sandy areas, and a 
small wetland. Each of these areas has a unique flora. The majority of the park 
is mesic upland woods, characterized by large, well-spaced trees, particularly 

Botany: Crandall and Dolan Vol. 106 (1997) 

Quercus alba, Acer saccharum, and Platanus occidentalism with very little veg- 
etation on the forest floor. A similar remnant Tipton Till Plain forest was 
reported by Rothrock, et al. (1994) in Mounds State Park. In locations where 
sunlight breaks through the trees, a burst of understory growth, often comprised 
of Cystopteris fragilis and Impatiens capensis, occurs. This burst of growth occurs 
most frequently at the edge of the woods where the greatest abundance of herba- 
ceous plants are found. Many areas of the woods host dramatic displays of spring 
ephemerals, such as Erigenia bulbosa, Mertensia virginica, Erythronium 
americanum, and Sanguinaria canadensis. 

The lowland sandy areas are located primarily along Crooked Creek, which 
divides the park into two nearly equal halves. The substrate in this area varies 
from recent sand deposits to sandy loam and supports a very different plant com- 
munity than the mesic upland woods. Such species as Justicia americana and 
Polygonum hydropiperoides were found in this area. Many of the plants in this 
habitat must be adapted to life in dry sand as well as in standing water due to the 
great fluctuations in the level of Crooked Creek throughout the growing season. 

The wetland is located in direct sunlight near the parking lot and is inhabit- 
ed by Elocharis ovata, Mimulus ringus, and Rumex crispus. The vegetation is 
prominent through July. After that time, the wetland dries up almost complete- 
ly and is eventually mowed. During years of high rainfall, the area may remain 
wet throughout the summer. 

A total of 2 fern and fern allies and 177 angiosperms have been identified at 
Crooked Creek Community Juan Soloman Park. Of the 179 taxa found, 126 
are native to the Midwest. Because 29.6% of the species were alien, the entire 
park cannot be a pristine natural area. However, the majority of the exotic plants 
were primarily if not exclusively located near the highly disturbed roadside. 
Away from this area, the flora is largely native with the exception of a few exot- 
ic invasives, such as garlic mustard (Alliaria petiolata) and wintercreeper (Euony- 
musfortunei). One plant collected, the lesser celendine (Ranunculus ficaria subsp. 
bulbifera), is a State record. Lesser celendine is an exotic species that has like- 
ly escaped from horticulture. 

When compared to other similar sites in the State of Indiana, Crooked Creek 
Community Juan Soloman Park has a large number of native species per acre. 
Crooked Creek has an average of 2.7 native plants per acre, compared to 2.0 
for the Fall Creek Nature Preserve in Warren County (Tonkovich and Sargent, 
1993) and 1.5 species per acre in Mounds State Park in Madison County (Rothrock, 
etaU 1993). 

The coefficient of conservatism (Swink and Wilhelm, 1994) was 54.1, indi- 
cating that Crooked Creek Community Juan Soloman Park is a high-quality nat- 
ural area in spite of the large number of non-native species present. The quality 
index values (C) are given in the description that follows each species' name in 
the checklist. Because exotic species have the potential to decrease the plant 
diversity (Bratton, 1982), Crooked Creek Community Juan Soloman Park needs 
a management plan in order to retain its quality. The invasive exotics of great- 

Vol. 106 (1997) Indiana Academy of Science 

est concern in the park are garlic mustard (Alliaria petiolata), wintercreeper 
(Euonymus fortunei), and amur bush honeysuckle {Lonicera maackii). 

Garlic mustard is most effectively controlled by pulling the immature plants, 
including their roots, from the ground and removing all pulled plants before seed- 
set (Nuzzo, etal, 1991). This process is most effective before the species becomes 
fully established in high-density populations. Fortunately, although garlic mus- 
tard is present in many areas in the park, its numbers are low at any given site. 

Wintercreeper is invading many areas of the mesic upland woods in the park 
both in the sun and shade. Eradication efforts should include cutting each vine 
by hand and spraying the plant with an herbicide that is non-toxic to aquatic 
organisms in the event that any runoff reaches Crooked Creek. Spraying 
should be done in the spring before the emergence of the spring ephemerals or 
during late autumn when most of the native plants are dormant (Hutchison, 1991). 
This practice must be continued in subsequent seasons to insure that all of the 
wintercreeper has been eliminated and that new invading individuals are not 
being introduced. 

Amur bush honeysuckles can tolerate many different habitat types and mois- 
ture regimes, so they are easily established and spread. The honeysuckles inhib- 
it the growth of native species through both shading and by releasing a growth-inhibiting 
chemical into the ground (Nyboer, 1991). To effectively eliminate amur bush 
honeysuckle, the plants should be cut and the stumps treated with herbicide 
(Nyboer, 1991). In addition, entire seedlings, including their roots, should be 
removed by hand-pulling when the soil is moist (Nyboer, 1991). 

Management and control of invasive exotics is crucial to the maintenance 
of the high quality habitats currently present in the park. Follow-up studies will 
be needed to determine whether the control efforts have been effective. Fur- 
thermore, the small wetland area identified in this study should be protected dur- 
ing any additional development activity in the park. 


The authors would like to thank the Butler Board of Trustees and the But- 
ler Grant Committee for providing the funding needed to make this learning 
experience possible. In addition, they would like to thank Kerry Manders of 
the Crooked Creek Community Council and Don Miller of the Indianapolis Parks 
Department for providing a collecting permit and approving the research. Final- 
ly, they would like to thank Mike Homoya and Helene Stares for providing assis- 
tance with plant identification and Dr. Andree Desrocheres for providing helpful 
comments on the manuscript. 


Bratton, S.P. 1982. The effects of exotic plant and animal species on nature preserves. Natur. Areas J. 2: 3- 

Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent 

Canada. New York Bot. Gard., New York, 910 pp. 

Botany: Crandall and Dolan Vol. 106 (1997) 

Herman, K.D., L.A. Master, M.R. Penskar, A.A. Reznicek, G.S. Wilhelm, and W.W. Brodowicz. 1997. Floris- 
tic quality assessment: Development and application in the State of Michigan. Natur. Areas J. 17: 265- 

Homoya, M.A., D.B. Abrell, J.R. Aldrich, and T.W. Post. 1985. The natural regions of Indiana. Proc. Indiana 
Acad. Sci. 94: 245-268. 

Hutchison, M. 1990. Vegetation management guidline: Wintercreeper or climbing euonymus (Euonymus 
fortunei). Illinois Nature Preserves Comm. Vegetation Manage. Manual 1(27): 4 pp. 

Indiana Department of Natural Resources. 1993. Indiana's rare plants and animals: A checklist of endan- 
gered and threatened species. Indianapolis, Indiana, 26 pp. 

Ladd, D.M. in prep. The Missouri floristic quality assessment system. The Nature Conservancy, St. Louis, 

Nuzzo, V, J. Kenney, and G. Fell. 1991. Vegetation management guidline: Garlic mustard (Allariaria petio- 
lata (Bieb.) Cavara & Grande). Illinois Nature Preserves Comm. Vegetation Manage. Manual 1(10): 5 

Nyboer, R. 1990. Vegetation management guidline: Bush honeysuckles: Tartarian, morrow's, belle, and armur 
honey suckle (Lonicera tatarica L., L. morrowii Gray, L. x bella Zabel, and L. maackii (Rupr.) Maxim). 
Illinois Nature Preserves Comm. Vegetation Manage. Manual 1(6): 4 pp. 

Rothrock, P.E., H. Stares, R. Dunbar, and R.L. Hedge. 1993. The vascular flora of Mounds State Park, Madi- 
son County, Indiana. Proc. Indiana Acad. Sci. 102: 161-199. 

Swink, F. and G. Wilhelm. 1994. Plants of the Chicago region. Indiana Acad. Sci., Indianapolis, Indiana, 921 

Tonkovich, G.S. and M.L. Sargent. 1993. The vascular plants of Fall Creek Gorge Nature Preserve, Warren 
County, Indiana. Proc. Indiana Acad. Sci. 102: 9-45. 

Sturm, R.H. and R.H. Gilbert. 1978. Soil survey of Marion County, Indiana. U.S. Dep. Agr. Soil Conserv. 
Serv., Washington D.C., 63 pp. 

Vol. 106 (1997) Indiana Academy of Science 





Aspleniaceae — Spleenwort Family 

Cystopteris fragilis (L.) Bernh. \a.r.fragilis 

Fragile fern; frequent near the border of Section 1 towards the 
soccer field; C = 10;CN = 81. 


Equisetaceae — Horsetail Family 

Equisetum arvense L. 

Common or field horsetail; occasional in sandy soil along the creek 
in Section 5; C = 0; CN = 168. 


Acanthaceae — Acanthus Family 

Justicia americana (L.) Vahl. 

Water- willow; common on the sandy banks of the creek in direct 
sunlight; C = 6; CN = 94. 

Ruellia strepens L. 

Smooth ruellia; one isolated population in Section 2 under a dog- 
wood across from the pavilion; C = 8; CN = 91. 

Aceraceae — Maple Family 

Acer negundo L. 

Boxelder or ash-leaved maple; common throughout the park; 
C = 0; CN = 48. 

Acer rubrum L. 

Red maple; occasional throughout the mesic upland woods in all 
sections; C = 7; not collected. 

Acer saccharum Marshall 

Sugar maple; common throughout the park; C = 3; CN = 47. 

C = The quality index of Swink and Wilhelm (1994). 

CN = Raelene Crandell's collection number. The Friesner Herbarium no longer 
assigns accession numbers. 

Botany: Crandall and Dolan Vol. 106 (1997) 

Anacardiaceae — Sumac Family 

Toxicodendron radicans (L.) Kuntze 

Poison ivy; abundant throughout the park, often as a vine; C = 2; 
not collected. 

Annonaceae — Custard Apple Family 

Asimina triloba (L.) Duna 

Pawpaw; frequent in the under story throughout all sections; C= 14; 

CN = 76. 

Apiaceae — Carrot Family 

Cryptotaenia canadensis (L.) DC. 

Honewort; common throughout the mesic upland woods of Sections 
1-4;C = 2;CN = 64. 

Daucus carota L. 

Wild carrot or Queen Anne's lace; one isolated population in Sec- 
tion 1 near Grandview Drive; CN =119 (alien). 

Erigenia bulbosa (Michx.) Nutt. 

Harbinger of spring; common in Sections 1 and 2 in both the low 
wetland areas and mesic upland woods; C = 10. 

Osmorhiza longistylis (Torr.) DC. 

Sweet cicely; common throughout the park in moist soil and near 
the border of the woods; C = 3; CN = 49. 

Sanicula marilandica L. 

Black snakeroot; common throughout the park; C = 6; CN = 73. 

Araceae — Arum Family 

Arisaema triphyllum (L.) Schott 

Jack in the pulpit; one isolated population in Section 3 under 
heavy tree cover; C = 4; not collected. 

Aristolochiaceae — Birthwort Family 

Asarum canadense L. 

Wild ginger; abundant in large clumps in the mesic upland areas of 
Sections 1-4; C = 7; CN = 24. 

Asclepiadaceae — Milkweed Family 

Apocynum cannabinum L. 

Indian hemp; occasional in the mesic upland woods of Sections 1 
and2;C = 4;CN=104. 

Asclepias incarnata L. 

Swamp milkweed; occasional in sand and direct sunlight near the 
creek in Section 5; C = 4; CN = 137. 

Vol. 106 (1997) Indiana Academy of Science 

Asclepias syriaca L. 

Common milkweed; occasional at the edge of the mesic upland 
woods in Sections 3 and 4; C = 0; CN = 1 14. 

Asteraceae — Aster Family 

Ambrosia artemisiifolia L. 

Common ragweed; abundant throughout the park in all sections; 
C = 0;CN=150. 

Ambrosia trifida L. 

Giant ragweed; occasional among Ambrosia artemisiifolia L. in all 
sections; C = 0; CN = 149. 

Aster ericoides L. 

Heath aster or squarrose white aster; common at the edge of the 
woods in Sections 1-4; C = 5; CN = 163. 

Bidens cernua L. 

Nodding bur-marigold; occasional in sandy soil along the stream in 
Sections 1, 2, and 5; C = 5; CN = 164. 

Cirsium arvense (L.) Scop. 

Canada thistle; abundant along the disturbed roadside in Sections 
l,3,and4;CN= 110 (alien). 

Erigeron annuus (L.) Pers. 

Daisy fleabane; common in sparsely wooded areas and near the 
border of the woods; C = 0; CN = 3 1 . 

Eupatorium maculatum L. 

Spotted joe-pye weed; occasional along the edge of the woods in 
Sections 1 and 2; C = 4; CN = 144. 

Eupatorium perfoliatum L. 

Boneset; occasional near the creek in sandy soil; C = 4; CN = 130. 

Eupatorium rugosum Houtt. 

White snakeroot; frequent in direct sunlight throughout all sec- 
tions of the park; C = 4; CN = 15 1 . 

Helenium autumnale L. 

Sneezeweed; occasional in sandy soil near the stream in Sections 1, 
2, and5;C=5;CN= 167. 

Helianthus divaricatus L. 

Woodland sunflower; common at the edge of the woods in all 
sections and in sand near the creek; C = 5; CN = 166. 

Heliopsis helianthoides (L.) Sweet. 

False sunflower; rare in sand and direct sunlight near the creek in 
Section 5; C = 5; CN = 124. 

10 Botany: Crandall and Dolan Vol. 106 (1997) 

Lactuca canadensis L. 

Tall lettuce; frequent at the edge of the woods in partial sunlight; 
C = 2; not collected. 

Lactuca floridana van villosa (L.) Gaertner 

Woodland or blue lettuce; abundant along the edge of the woods in 
partial sunlight; C = 5; CN = 134. 

Lactuca serriola L. 

Prickly lettuce; isolated population located along the disturbed 
roadside in Section 1; CN =111 (alien). 

Polymnia canadensis L. 

Pale-flowered leaf-cup; abundant at the edge of the woods through- 
out all sections; C = 10; CN = 146. 

Rudbeckia fulgida Ait. 

Eastern coneflower; frequent at the edge of the woods in partial 
sunlight; C = 8; CN = 135. 

Senecio jacobaea L. 

Tansy-ragwort; rare, one plant in Section 2 at the edge of the woods; 
not collected (alien). 

Silphium perfoliatum L. 

Cup-plant; one population in Section 5 in sand and direct sunlight 
near the creek; C = 5; CN = 139. 

Solidago canadensis L. 

Common goldenrod; occasional throughout the park and near the 
creek in sunny areas; C = 1; CN = 164. 

Sonchus oleraceus L. 

Common sow-thistle; common along the roadside in Sections 3 and 
4; CN= 131 (alien). 

Taraxacum officinale Weber ex Wiggers 

Common dandelion; common in direct sunlight and open areas 
among grass; CN = 35 (alien). 

Verbesina alternifolia (L.) Britton 

Wingstem; common throughout the park in sunny areas; C = 5; 
CN = 147. 

Balsaminiaceae — Jewel-Weed Family 

Impatiens capensis Meerb. 

Spotted touch-me-not; abundant in mesic upland areas among 
stinging nettle; C = 3; CN = 71. 

Vol. 106 (1997) Indiana Academy of Science 1 1 

Impatiens pallida Nutt. 

Pale or yellow touch-me-not; occasional in mesic upland areas, 
not as abundant as Impatiens capensis Meerb.; C = 6; CN = 83. 

Berberidaceae — Barberry Family 

Podophyllum peltatum L. 

May apple; frequent throughout all mesic upland areas, often 
occurring in large clumps; C = 4; CN = 21. 

Betulaceae — Birch Family 

Ostrya virginiana (Miller) K. Koch. 

Hop-hombeam or ironwood; occasional throughout the mesic upland 
woods of all sections; C = 5; not collected. 

Boraginaceae — Borage Family 

Mertensia virginica (L.) Pers. 

Eastern bluebell; rare, in sandy soil near the creek in Section 2; 
C = 5;CN=11. 

Brassicaceae — Mustard Family 

Alliaria petiolata (Bieb.) Cavara & Grande 

Garlic-mustard; exotic invasive, abundant throughout all sections; 
CN= 19 (alien). 

Barbarea vulgaris R. Br. 

Yellow rocket; occasional in direct sunlight in Sections 1 and 2 
among the mowed grass; CN = 97 (alien). 

Cardamine concatenata (Michx.) O. Schwarz 

Five-parted toothwort; common in the mesic upland areas of 
Sections 1-4 where the trees are sparse; C = 5; CN = 3. 

Cardamine douglassi Britton 

Purple cress; isolated population in the open wetland area of 
Section 1 near the parking lot; C = 7; CN = 70. 

Hesperis matronalis L. 

Dame's rocket; common throughout the mesic upland woods of 
Sections 1-4, particularily at the edges; CN = 30 (alien). 

Lepidium virginicum L. 

Poor-man's-pepper or pepper-grass; rare, one small, isolated 
population in the mesic upland woods of Section 1 ; CN = 86 (alien). 

Caesalpiniaceae — Caesalpinia Family 

Cercis canadensis L. 

Redbud; occasional in mesic upland areas; C = 10; CN = 72. 

12 Botany: Crandall and Dolan Vol. 106 (1997) 

Gleditsia triacanthos L. 

Honey-locust; occasional throughout the park in all sections; C = 2; 
not collected. 

Gymnocladus dioica (L.) K. Koch. 

Kentucky coffee-tree; occasional in mesic upland areas; C = 8; not 

Campanulaceae — Bellflower Family 

Campanula americana L. 

American bellflower; one population in a small clearing in sandy 
soil;C = 3;CN= 126. 

Lobelia siphilitica L. 

Great lobelia; occasional in sand near the creek in Sections 1 and 2; 
C = 6;CN=158. 

Caprifoliaceae — Honeysuckle Family 

Lonicera japonica Thunb. 

Japanese honeysuckle; isolated population in Section 2 climbing on 
some trees and shrubs; CN = 77 (alien). 

Lonicera maackii (Rupr.) Maxim. 

Amur bush honeysuckle; common in Sections 1-4 along the edge 
of the woods; CN = 60 (alien). 

Lonicera oblongifolia (Goldie) Hook 

Swamp fly honeysuckle; occasional in mesic upland areas; C = 10; 

CN = 28. 

Caryophyllaceae — Pink Family 

Saponaria officinalis L. 

Bouncing bet; one isolated population in sand and direct sunlight 
near the creek in Section 5; CN =121 (alien). 

Silene nivea (Nutt.) Otth. 

White campion; common in sandy soil near the creek; C = 10; 

CN = 90. 

Stellaria media (L.) Villars. 

Common chickweed; common along the border of the woods in 
Sections 1 and 2; CN = 33 (alien). 

Celastraceae — Bittersweet Family 

Celastrus scandens L. 

American bittersweet; occasional throughout the mesic upland woods 
of all sections; C = 4; not collected. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 1 3 

Euonymus atropurpureus Jacqs. 

Wahoo; one shrub at the border of the woods in Section 2 that may 
have been planted for horticultural reasons; C = 8; CN = 78. 

Euonymus fortunei (Turcz.) Hand.-Mazz. 

Wintercreeper; exotic invasive throughout mesic upland woods of 
Sections 1-4; CN = 105 (alien). 

Chenopodiaceae — Goosefoot Family 

Chenopodium album L. 

Lamb's quarters or pigweed; occasional along the disturbed road- 
side in Sections 3 and 4; CN =132 (alien). 

Clusiaceae — Mangosteen Family 

Hypericum mutilum L. 

Dwarf St. John's-wort; occasional near the creek in sandy soil and 
partial sunlight in Section 5; C = 8; CN = 118. 

Commelinaceae — Spiderwort Family 

Commelina communis L. 

Common day-flower; occasional in sand near the creek in Section 
5; CN= 136 (alien). 

Tradescantia virginiana L. 

Spiderwort; occasional in the mesic upland woods of Sections 1-4; 
C = 5;CN=102. 

Convolvulaceae — Morning-glory Family 

Calystegia sepium (L.) R.Br. 

Hedge-bindweed; locally abundant in Section 5 along the creek in 
sand and direct sunlight; C = 1; CN =113. 

Cornaceae — Dogwood Family 

Cornus drummondii C.A. Meyer 

Rough-leaved dogwood; occasional in Sections 1 and 2, most often 
near the border of the woods; C = 2; CN = 75. 

Cucurbitaceae — Gourd Family 

Sicyos angulatus L. 

Bur-cucumber; occasional in Sections 1 and 2 climbing on small 
shrubs; C = 5; CN = 157. 

Cyperaceae — Sedge Family 

Carex amphibola Steudel 

Sedge; common throughout the mesic upland woods of Sections 
1-4;C=10;CN = 58. 

14 Botany: Crandall and Dolan Vol. 106 (1997) 

Carex davisii Schwein & Torr. 

Sedge; occasional in sandy soil under heavy tree cover in Sections 
l,2,and5;C = 7;CN = 87. 

Carex grayi Carey 

Sedge; occasional in moist, sandy soil; C = 7; CN =127. 

Carex shortiana Dewey 

Sedge; one isolated population in sandy soil on the trail through 
Section 1;C= 10;CN = 89. 

Elocharis ovata (Roth) Roemer & Schultes 

Blunt spike-rush; locally abundant in the open wetland in Section 
1 near the parking lot; C = 10; CN = 66. 

Dipsacaceae — Teasel Family 

Dipsacus sylvestris Huds. 

Common teasel; abundant along the disturbed roadside in Sections 
1,3, and 4; CN = 112 (alien). 

Fabaceae — Pea Family 

Melilotus officinalis (L.) Pallas 

Yellow sweet clover; one isolated population in Section 4 near Fox 
Hill Road; CN = 99 (alien). 

Robinia pseudoacacia L. 

Black locust; common throughout the mesic upland woods; 

CN = 42. 

Trifolium dubium Sibth. 

Little hop-clover; frequent in direct sunlight in the open grassy areas 
of Sections 1 and 2; CN = 37 (alien). 

Trifolium repens L. 

White clover; abundant in direct sunlight in the open grassy areas 
of Sections 1 and 2; CN = 36 (alien). 

Fagaceae — Beech Family 

Fagus grandifolia Ehrh. 

American beech; frequent throughout the park; C = 5; CN = 122. 

Quercus alba L. 

White oak; occasional throughout the mesic upland woods of all 
sections; C = 5; not collected. 

Quercus muhlenbergii Engelm. 

Yellow oak; common in the mesic upland woods of Sections 1-4; 
C = 8; CN = 84. 

Vol. 106(1997 Indiana Academy of Science 15 

Quercus palustris Muenchh. 

Pin-oak; occasional throughout the mesic upland woods of all 
sections; C = 8; not collected; likely planted. 

Quercus rubra L. 

Red oak; the edge of the woods near the pavilion; not as common 
as the other oaks; C = 7; CN = 84. 

Quercus velutina Lam. 

Black oak; occasional throughout the mesic upland woods of all 
sections; C = 6; not collected. 

Fumariaceae — Fumitory Family 

Dicentra cucullaria (L.) Bernh. 

Dutchman's breeches; frequent throughout the mesic upland woods 
of Sections 1-4; C = 6; CN = 2. 

Grossulariaceae — Gooseberry Family 

Hydrangea paniculata Siebold 

Hydrangea; occasional in Section 5 in sandy soil and partial 
sunlight; CN = 129 (alien). 

Hydrophylaceae — Waterleaf Family 

Hydrophyllum appendiculatum Michx. 

Biennial waterleaf; occasional in sandy soil and throughout the mesic 
upland woods of Sections 1-4; C = 8; CN = 26. 

Iridaceae — Iris Family 

Sisyrinchium angustifolium Miller 

Blue-eyed grass; one isolated population in Section 1 at the edge of 
the woods near Grandview Drive; C = 10; CN = 34. 

Juglandaceae — Walnut Family 

Carya cordiformis (Wangenh.) K. Koch. 

Bitternut hickory; occasional throughout the mesic upland woods 
of all sections; C = 7; not collected. 

Carya ovata (Miller) K. Koch. 

Shagbark hickory; occasional in the mesic upland woods; C = 5; not 

Lamiaceae — Mint Family 

Glechoma hederacea L. 

Gill-over-the-ground; abundant throughout all sections; CN = 8 

16 Botany: Crandall and Dolan Vol. 106 (1997) 

Lamium amplexicaule L. 

Henbit; common in direct sunlight and at the edge of the woods in 
Sections 1 and 2; CN = 14 (alien). 

Leonurus cardiaca L. 

Motherwort; common near the edge of the woods near Fox Hill Road 
in Sections 3 and 4; CN = 101 (alien). 

Mentha arvensis L. 

Field mint; occasional in sandy soil near the creek; C = 5; CN =138. 

Prunella vulgaris L. 

Self-heal; occasional at the edge of the woods in Sections 1 and 2; 
CN = 98 (alien). 

Stachys tenuifolia Willd. 

Smooth hedge-nettle; occasional to frequent at the edge of the woods 
in Sections 1 and 2; C = 8; CN = 148. 

Teucrium canadense L. 

Germander; occasional in Section 1 at the edge of the woods; 
C = 3;CN=120. 

Liliaceae — Lily Family 

Allium tricoccum Aiton 

Ramps, wild leek; one small isolated population under heavy tree 
cover along the trail through Section 1; C = 7; CN = 93. 

Allium vineale L. 

Field-garlic; common throughout the park in partial to direct 
sunlight, especially near its borders; CN = 59 (alien). 

Camassia scilloides (Raf.) Cory 

Wild hyacinth; occasional in sandy soil in Sections 1 and 2; C = 6; 

CN = 10. 

Erythronium americanum Ker-Gawl. 

Yellow trout lily; common in the heavily shaded areas of the mesic 
upland woods of Sections 1-4; C = 8; CN = 5. 

Hemerocallis fulva (L.) L. 

Day-lily; common along the disturbed roadside in Sections 3 and 4 
near Fox Hill Road; CN = 103 (alien). 

Narcissus pseudo-narcissus L. 

Daffodil; two isolated plants in sandy soil near the creek; not 
collected (alien). 

Ornithogalum umbellatum L. 

Star of Bethlehem; rare, two isolated populations in Sections 1 and 
3; not collected (alien). 

Vol. 1 06 ( 1 997) Indiana Academy of Science 1 7 

Polygonatum biflorum (Walter) Elliott 

Soloman's seal; abundant in sandy soil and throughout the mesic 
upland woods of all sections; C = 3; CN = 32. 

Trillium recurvatum Beck 

Red trillium; common in the heavily shaded areas of the mesic upland 
woods of Sections 1-4; C = 5; CN = 18. 

Moraceae — Mulberry Family 

Morus alba L. 

White mulberry; frequent throughout all sections; CN = 51 (alien). 

Oleaceae — Olive Family 

Fraxinus americana L. 

White ash; common throughout the mesic upland woods of Sections 
1-4;C = 5;CN = 50. 

Fraxinus pennsylvanica Marshall 

Green ash; occasional along the creek in Section 5 and rare through- 
out the mesic upland woods of Sections 1-4; C = 5; not collected. 

Onagraceae — Evening Primrose Family 

Circaea lutetiana L. 

Enchanter's nightshade; abundant in Sections 1 and 2; C = 1; 

CN = 82. 

Oenothera biennis L. 

Common evening-primrose; rare along the disturbed roadside in 
Sections 1 and 3; C = 0; not collected. 

Oxalidaceae — Wood Sorrel Family 

Oxalis grandis Small 

Yellow wood-sorrel; abundant in direct sunlight in the open grassy 
areas of Sections 1 and 2; CN = 38 (alien). 

Papaveraceae — Poppy Family 

Sanguinaria canadensis L. 

Bloodroot; common throughout the mesic upland woods of Sections 
1-4;C = 6;CN=1. 

Stylophorum diphyllum (Michx.) Nutt. 

Celandine-poppy; one isolated population in Section 2 across from 
the playground in a sparsely wooded area; C = 10; CN = 27. 

Phytolaccaceae — Pokeweed Family 

Phytolacca americana L. 

Pokeweed; two isolated populations in Sections 1 and 4: C = 1; 

18 Botany: Crandall and Dolan Vol. 106 (1997) 

Plantaginaceae — Plantain Family 

Plantago lanceolata L. 

English plantain; common near the disturbed roadside in Sections 
3and4;CN = 85, 160 (alien). 

Plantago major L. 

Common plantain; common in the open areas of Sections 1 and 2 
among the mowed grass; CN =106 (alien). 

Platanaceae — Plane-Tree Family 

Platanus occidentalis L. 

Sycamore; common throughout all sections; C = 6; not collected. 

Poaceae — Grass Family 

Dactylis glomerata L. 

Orchard grass; common throughout all sections; CN = 57 (alien). 

Eleusine indica (L.) Gaertn. 

Yard grass; common throughout the park in the open, disturbed areas 
of all sections; CN = 140 (alien). 

Elymus riparius Wieg. 

Streambank wild rye; common throughout the mesic upland woods 
of Sections 1-4; C = 5; CN = 107. 

Elymus villosus Muhl. 

Downy wild rye; common in sandy soil in Sections 1, 2, and 5; 
C = 5; CN = 65 (alien). 

Elytrigia repens (L.) Nevski. 

Quack-grass; common in the open areas of Sections 1 and 2; 
CN = 62 (alien). 

Festuca rubra L. 

Red fescue; frequent throughout all sections of the park; CN = 56 

Phalaris arundinacea L. 

Reed canary grass; frequent at the edge of the creek in sandy soil 
in Sections 1, 2, and 5; CN = 88 (alien). 

Phleum pratense L. 

Timothy; common throughout the mesic upland woods of Sections 
1-4; CN = 108 (alien). 

Poa pratensis L. 

Kentucky bluegrass; frequent throughout all sections of the park; 
CN = 61 (alien). 

Vol. 1 06 ( 1 997) Indiana Academy of Science 1 9 

Setaria glauca (L.) P. Beauv. 

Yellow foxtail grass; frequent near the edge of Fox Hill Road and 
in other disturbed areas in Sections 1,3, and 4; CN = 159, 161 (alien). 

Polemoniaceae — Phlox Family 

Phlox divaricata L. 

Common phlox; occasional in sandy soil in Sections 1 and 2; 
C = 5;CN=13. 

Phlox paniculata L. 

Summer phlox; occasional in sandy soil along the creek; CN = 143. 

Polygonaceae — Buckwheat Family 

Polygonum amphibium L. 

Water smartweed; occasional in sand near the creek in Sections 1, 
2, and 5 among Polygonum lapathifolium L.; C = 4; CN = 154. 

Polygonum hydropiperoides Michx. 

False water-pepper; occasional in moist sand and often standing 
water near the creek in Section 5; C = 7; CN =128. 

Polygonum lapathifolium L. 

Pale smartweed; occasional in sand near creek in Sections 1, 2, 
and5;C = 0;CN=155. 

Polygonum pensylvanicum L. 

Smartweed; common in disturbed areas near the roadside and near 
the border of the woods in Sections 1-4; C = 0; CN = 68. 

Polygonum persicaria L. 

Lady's thumb; common along Fox Hill Road in Sections 3 and 4; 
CN = 141 (alien). 

Polygonum scandens L. 

False buckwheat; occasional in all sections of the park climbing 
on small shrubs; C = 1; CN = 152. 

Polygonum virginianum L. 

Virginia knotweed or jumpseed; occasional throughout the park in 
partial sunlight; C = 2; CN = 156. 

Rumex crispus L. 

Curly dock; common in the open wetland in Section 1 and in the 
mesic upland woods of Sections 1-4; CN = 67 (alien). 

Pontederiaceae — Water-Hyacinth Family 

Pontederia cordata L. 

Pickerel- weed; one plant found in the creek along the edge of the 
trail in Section 1; C = 10; not collected. 

20 Botany: Crandall and Dolan Vol. 106 (1997) 

Portulaceae — Purslane Family 

Claytonia virginica L. 

Spring-beauty; frequent in direct sunlight and at the edge of the 
woods in Sections 1-4; C = 2; CN = 4. 

Portulaca oleracea L. 

Common purslane; common growing out of the cracks in the asphalt 
near the edge of the parking lot in Section 1; CN = 142 (alien). 

Primulaceae — Primrose Family 

Lysimachia ciliata L. 

Fringed loosestrife; common in Sections 1 and 2 in partial to direct 
sunlight; C = 4; CN = 79. 

Ranunculaceae — Buttercup Family 

Anemonella thalictroides (L.) Spach. 

Rue anemone; common throughout Sections 1 and 2; C = 7; 

Ranunculus abortivus L. 

Small-flowered crowfoot; common in sandy soil and the mesic 
upland woods of Sections 1-4; C = 0; CN = 20. 

Ranunculus ficaria L. subsp. bulbifera Lambionon 

Lesser celandine; one isolated population in sandy soil in Section 
2; CN = 7 (alien). 

Rosaceae — Rose Family 

Agrimonia gryposepala Wallr. 

Common agrimony; occasional at the edge of the woods in partial 
sunlight in Section 2; C = 2; CN = 145. 

Crataegus sp. L. 

Hawthorn; common in the mesic upland woods; C = 4; not 

Geum canadense Jacq. 

White avens; only a couple dozen plants along the edge of the woods 
in Sections 1 and 2; C = 1; CN = 80. 

Prunus serotina Ehrh. 

Wild black cherry; common throughout the mesic upland woods 
of Sections 1-5; C = 1 ; CN = 43. 

Rosa Carolina L. 

Pasture rose; one isolated population in Section 4 near Fox Hill 
Road.;C = 5;CN= 100. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 2 1 

Rosa multiflora Thunb. 

Multiflora rose; one shrub in Section 2 at the edge of the woods that 
may have been planted for horticultural reasons; CN = 39 (alien). 

Rubus allegheniensis T.C. Porter 

Common blackberry; one shrub in Section 2 at the edge of the woods 
that may have been planted for horticultural reasons; C = 3; 

CN = 40. 

Rubiaceae — Madder Family 

Gallium aparine L. 

Cleavers; abundant in all mesic upland areas where myrtle is not 
abundant; C= 1;CN = 23. 

Salicaceae — Willow Family 

Populus deltoides Marshall 

Common cottonwood; occasional in the mesic upland areas; 

C = 2;CN = 52. 

Salix babylonica L. 

Weeping willow; one tree in Section 3 near the bridge on Grand- 
view Drive; C = 53 (alien). 

Salix nigra Marshall 

Black willow; occasional throughout the mesic upland woods of 
Sections 3 and 4; C = 4; not collected. 

Scrophulariaceae — Figwort Family 

Mimulus ring ens L. 

Monkey-flower; common in direct sunlight in Section 1 among 
the mowed grass and in the wetland; C = 6; CN = 96. 

Verbascum thapsus L. 

Common mullein; occasional along the roadside in Sections 3 and 
4; CN= 123 (alien). 

Veronica arvensis L. 

Corn speedwell; common in Sections 1 and 2 in direct sunlight 
among the grass; often occurs with Lamium amplexicaule L.; 
CN = 15 (alien). 

Veronica peregrina L. 

Purslane speedwell; common in Sections 1 and 2 in direct sunlight 
among the grass; C = 0; CN = 29. 

Simaroubaceae — Quassia Family 

Ailanthus altissima (Miller) Swingle 

Tree of heaven; only one isolated population in Section 1, but 
common in Sections 3-4; CN = 95 (alien). 

22 Botany: Crandall and Dolan Vol. 106 (1997) 

Solanaceae — Nightshade Family 

Solarium carolinense L. 

Horse-nettle; occasional to rare in Sections 1 and 4 near the edge of 
the woods; CN = 109 (alien). 

Solarium dulcamara L. 

Bittersweet; common in Sections 1-3 in partial to direct sunlight; 
CN = 54 (alien). 

Tiliaceae — Linden Family 

Tilia americana L. 

Basswood or American linden; occasional throughout the park; 
C = 5; not collected. 

Ulmaceae — Elm Family 

Celtis occidentalis L. 

Northern hackberry; common throughout the park; C = 3; not 

Ulmus rubra Muhl. 

Red or slippery elm; common throughout the mesic upland woods 
of Sections 1-5; C = 4; CN = 46. 

Urticaceae — Nettle Family 

Pilea pumila (L.) Gray 

Clearweed; common along the edge of the woods in direct 
sunlight in Sections 1 and 2; C = 5; CN = 153. 

Urtica dioica L. 

Stinging nettle; abundant in the mesic upland woods and through- 
out all sections, especially among jewelweed; not collected (alien). 

Verbenaceae — Vervain Family 

Phryma leptostachya L. 

Lopseed; common at the border of the mesic upland woods in 
Sections 1 and 2; C = 4; CN = 92. 

Verbena urticifolia L. 

White vervain; occasional at the border of the mesic upland woods 
in Sections 1 and 2; C = 5; CN = 116. 

Violaceae — Violet Family 

Viola cucullata Aiton. 

Blue marsh-violet; abundant throughout all sections; C = 9; 
CN = 6. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 23 

Viola pubescens Aiton. 

Yellow forest violet; common throughout Sections 1-4; C = 5; 

Viola striata Aiton. 

Cream or pale violet; abundant throughout Sections 1-4; C = 6; 

Vitaceae — Grape Family 

Parthenocissus quinquefolia (L.) Planchon 

Virginia-creeper; abundant in all sections; C = 2; CN = 49. 

Vitis vulpina L. 

Winter grape; abundant in all sections and common climbing on 
small shrubs; C = 9; CN = 125. 

Proceedings of the Indiana Academy of Science Z J 

(1997) Volume 106 p. 25-31 


Richard D. Hyerczyk 
5204 South Natoma Avenue 
Chicago, Illinois 60603-1222 

ABSTRACT: Twenty-nine species of lichens are reported from Hoosier Prairie 
State Nature Preserve in Lake County, Indiana. Thirteen are of the crustose 
growth form, thirteen are foliose, and three are fruticose. An annotated species 
list with information on the habitats and distribution for each species is pro- 
vided. The results of this study indicate that lichenized fungi are relatively 
uncommon at this nature preserve. 

KEYWORDS: Arenicolous, corticolous, crustose, foliose, fruticose, lichen, 
lignicolous, saxicolous. 


The Hoosier Prairie State Nature Preserve is located about 43 km (27 miles) 
southeast of downtown Chicago in Lake County, Indiana (Figure 1). This 177 
hectare (439 acre) State Nature Preserve, which is located in the Northwestern 
Morainal Natural Region (Homoya, et al., 1985), is owned and managed by 
the Indiana Department of Natural Resources. The prairie developed on beach 
sand lying over lake bottom clay after Lake Michigan water levels receded from 
this area about 9,000 years ago. 

The topography is nearly level, with the elevation averaging 189 m (620 
feet). Soils are generally sandy and predominantly of the Brems, Maumee, and 
Watseka Series (Persinger, 1972). For northwestern Indiana, the average Janu- 
ary temperature ranges from a high of -1° C (30° F) to a low of -9°C (16° F), and 
the average July temperature, from a high of 28° C (83° F) to a low of 17° C (63° 
F). An average of 91 cm (36") of precipitation falls per year (Bair, 1992). 

Several natural communities are found at Hoosier Prairie. Dry savannas 
occur on sand rises and are dominated by Quercus alba L. and Q. velutina Lam. 
Mesic sand prairie openings lie between the rises and swales and are dominat- 
ed by Populus deltoides Marshall, P. tremuloides Michx., and Salix interior 
Rowlee. Wet prairies, sedge meadows, and marshes are scattered throughout the 
preserve in depressions and flats. 

Management of the prairie includes brush cutting and prescribed burning. 
Other human influences include concrete curbing in a gravel parking lot, wood 
rail fencing, a mowed hiking trail, and a few piles of concrete rubble. The sur- 
rounding land is approximately 75% residential and 25% industrial. 

Hoosier Prairie is within the boundaries of Calkins' (1896) flora, but none 
of the 125 species he reported were cited specifically from Lake County. Indi- 
ana. Wetmore (1986) reported 62 species of lichens from the Indiana Dunes 
National Lakeshore, but he did not include any from nearby Hoosier Prairie. 


Botany: Hyerczyk 

Vol. 106 (1997) 

Main St. 

Hoosier Prairie State Nature Preserve 

Map courtesy of the Indiana Department of Natural Resources: 
Division of Nature Preserves 

Figure 1 . Hoosier Prairie Nature Preserve. 

Since no lichenological studies were conducted in Hoosier Prairie, the purpose 
of this study was to provide information on the habitats and distribution of the 
lichen flora there. 

BetweenAugust 1991 and April 1997, six trips were made to Hoosier Prairie 
to collect voucher specimens and information on the habitats of these lichens. 

Vol. 106 (1997) Indiana Academy of Science 27 

Spot tests for chemical substances were made on the collected specimens using 
sodium hypochlorite and potassium hydroxide. Thin-layer chromatography (Cul- 
berson, 1972) was used to verify secondary-product chemistry of the Cladoniae. 
Specimens were identified using keys by Brodo (1988), Hale (1979), and Wil- 
helm (1995). A set of voucher specimens has been deposited in the herbariums 
at the Indiana Dunes National Lakeshore, Porter, Indiana, and at the Morton 
Arboretum, Lisle, Illinois. 


Twenty-nine species of lichenized fungi in 19 genera are reported from 
Hoosier Prairie State Nature Preserve (see Checklist). Thirteen species are of the 
crustose growth form, thirteen are foliose, and three are fruticose. Two species 
were common, three were frequent, ten were occasional, and fourteen were rare. 
The most common lichens were Candelaria concolor and Physcia millegrana. 
These two species are ubiquitous throughout northwestern Indiana. 

Nearly 62% of the flora was generally found on corticolous substrates (Quer- 
cus, Populus, and Salix spp.), 17% was lignicolous (on a wooden fence around 
the parking lot and on decorticate logs), 14% was saxicolous (on concrete 
curbing and rubble), and 7% was arenicolous (on sandy soil). 

Only 1 1 of the 125 species reported by Calkins were found at Hoosier Prairie. 
Of the 62 species reported by Wetmore, only 17 were found. The study areas of 
both Calkins and Wetmore were much larger than Hoosier Prairie and probably 
had more habitats and subtrates available for lichen colonization. Eleven species 
of lichens were found that were not included in Calkins' and Wetmore's studies. 
At least 6 of those species were found on substrates that were brought in for con- 
struction of the parking lot (concrete and wood) and were not found in a natur- 
al setting. 

The effect prescribed burning is having on the Hoosier Prairie lichen flora 
is not known, and no attempt was made to determine this. However, Wetmore 
(1981) mentions that frequent burning reduces lichen abundance, which may 
account for their low numbers here. 


The author would like to thank Dr. Gerould Wilhelm and Linda Masters, 
both of Conservation Design Forum, Inc., Elmhurst, Illinois, for doing the 
thin-layer chromatography on the Cladoniae and for their help and assistance in 
the identification and verification of specimens. Also, thanks to Tom Post, Region- 
al Ecologist with the Indiana Department of Natural Resources, and John A. 
Bacone, Director of the Indiana Division of Natural Resources, for permission 
to do this study at an Indiana State Nature Preserve and to two anonymous review- 
ers for their comments and suggestions. Finally, thanks to volunteer Deb Petro 
for initially suggesting a lichen study at Hoosier Prairie and for showing me 
around the Nature Preserve. 

28 Botany: Hyerczyk Vol. 106 (1997) 


Bair, F.E. 1992. The weather almanac, 6th ed. Gale Research, Inc., Detroit, Michigan, 855 pp. 

Brodo, I.M. 1988. Lichens of the Ottawa region, 2nd ed. Nat. Mus. Natur. Sci., Ottawa, Canada, 115 pp. 

Calkins, W.W. 1896. The lichen flora of Chicago and vicinity. Chicago Acad. Bull. 1, 50 pp. 

Culberson, C.F. 1972. Improved conditions and new data for the identification of lichen products by a stan- 
dardized thin-layer chromatography method. J. Chromatogr. 72: 113-125. 

Esslinger, T and R.C. Egan. 1995. A sixth checklist of the lichen-forming, lichenicolous and allied fungi of 
the continental United States and Canada. The Bryologist 98(4): 467-549. 

Hale, M.E., Jr. 1979. How to know the lichens, 2nd ed. Wm. C. Brown Co., Dubuque, Iowa, 246. pp. 

Homoya, M.A., D.B. Abrell, J.R. Aldrich, and T.W. Post. 1985. The natural regions of Indiana. Proc. Indiana 
Acad. Sci. 94: 245-268. 

Persinger, I.D. 1972. Soil survey of Lake County, Indiana. Purdue Univ. Agr. Exp. Station. U.S. Dep. Agr. Soil 
Conser. Serv., 66 pp. 

Wetmore, CM. 1981. Lichen studies on Allison Savanna. J. Minnesota Acad. Sci. 47: 2-3. 

. 1986. Lichens and air quality in the Indiana Dunes National Lakeshore. Mycotaxon 33: 25-39. 

Wilhelm, G. 1995. Lichens of the Chicago region. Unpub. manuscript, 95 pp. 

Vol. 106 (1997) Indiana Academy of Science 29 


The following is an annotated list of the lichenized fungi collected at Hoosier 
Prairie. Their arrangement is alphabetical by genus and then species. Presence, 
along with a brief description of habitat, is followed by the growth form and sub- 
strate(s), which are listed in brackets. The collection number is given in paren- 
theses. All collections were made by the author. Nomenclature and authority 
follow Esslinger and Egan (1995). Lichens reported by Wetmore (1986) are indi- 
cated by a "W." 

Amandinea Choisy ex Scheid. & H. Mayrh. 

Amandinea punctata (Hoffm.) Coppins & Scheid. Rare; on a weathered wood 
rail fence (196). [crustose/lignicolousj. W. 

Anisomeridium (Mull. Arg.) Choisy 

Anisomeridium nyssigenum (Ellis & Everh.) R.C. Harris. Rare; on the lower 
trunk of Quercus alba in a shaded mesic woodland (1371). The conidi- 
al state of this lichen, which has been called Sarcinulella banksiae Sut- 
ton & Alcorn, is represented here. [CRUSTose/corticolous]. 

Arthonia Ach. 

Arthonia caesia (Flotow) Korber. Occasional; on the trunks of Populus 
tremuloides and Quercus velutina (201). [CRUSTose/corticolous]. W. 

Caloplaca Th. Fr. 

Caloplacaferacissima H. Magn. Rare; on weathered concrete curbing (250). 


Caloplaca holocarpa (Hoffm. ex Ach.) M. Wade. Occasional; on a weath- 
ered wood rail fence and a decorticate log (350). [CRUSTOSE/lignicolous]. 

Caloplaca microphyllina (Tuck.) Hasse. Rare; on a weathered wood rail 
fence (193). [crustose/lignicolous]. 

Candelaria A. Massal. 

Candelaria concolor (Dickson) Stein. Common; on the lower branches of 
Populus deltoides and Quercus velutina (206). [foliose/corticolousj. 

Candelaria concolor var. effusa (Tuck.) G. Merr & Burnham. Occasional; 
on the trunks of Populus deltoides and Quercus velutina (813). [foliose 


30 Botany: Hyerczyk Vol. 106 (1997) 

Candelariella Mull. Arg. 

Candelariella reflexa (Nyl.) Lettau. Occasional; on a weathered wood rail 
fence and lower branches of Quercus velutina (1368). [CRUSTOSE / cor- 


Cladonia P. Browne 

Cladonia peziziformis (With.) J.R. Laundon. Rare; on sandy soil along a 
mowed hiking trail (214). [fruticose/arenicolous]. W. 

Cladonia polycarpoides Nyl. Rare; on sandy soil in a wet depression with 
Drosera intermedia (346). [fruticose/arenicolous]. W. 

Cladonia ramulosa (With.) J.R. Laundon. Occasional; at the base of Quer- 
cus velutina (1372). [fruticose/corticolousj. W. 

Cyphelium Ach. 

Cyphelium tigillare (Ach.) Ach. Rare; on a weathered wood rail fence (1494). 


Endocarpon Hedwig 

Endocarpon pusillum Hedwig. Rare; on concrete rubble (812). [CRUSTose/ 


Flavopunctelia (Krog) Hale 

Flavopunctelia flaventior (Stirton) Hale. Rare; on the lower trunk of 
Quercus rubra (205-A). [foliose/corticolousj. 

Hyperphyscia Mull. Arg. 

Hyperphyscia adglutinata (Florke) H. Mayrh. & Poelt. Occasional; on a 
decorticate oak log and on the lower branches of Quercus velutina (1318, 
1370) and Salix interior (1373). [foliose / corticolous - lignicolousj. 
Lecanora Ach. 

Lecanora dispersa (Pers.) Sommerf. Rare; on weathered concrete curbing 


Lecanora strobilina (Sprengel) Kieffer. Rare; on the lower trunk of Quer- 
cus alba (1495). [crustose/ corticolous]. 

Lecanora symmicta (Ach.) Ach. Frequent; on the lower trunks of Quercus 
velutina (202) and Salix interior and on a weathered wood rail fence 


Melanelia Essl. 

Melanelia subaurifera (Nyl.) Essl. Rare; on the lower branches of Salix inte- 
rior (1496). [FOLIOSE /CORTICOLOUS]. 

Vol. 106 (1997) Indiana Academy of Science 31 

Parmelia Ach. 

Parmelia sulcata Taylor. Frequent; on the lower branches of Populus tremu- 
loides (224), Quercus velutina, and Salix interior (207). [foliose/cor- 


Phaeophyscia Moberg 

Phaeophyscia pusilloides (Zahlbr.) Essl. Occasional; at the base of Quercus 
velutina (815) [foliose/corticolous]. W. 

Phaeophyscia rubropulchra (Degel.) Essl. Occasional; at the bases of Pop- 
ulus deltoides (210) and Salix interior (1369). [foliose/corticolous]. 

Physcia (Schreber) Michaux 

Physcia adscendens (Fr.) H. Olivier. Occasional; on the lower trunk of Pop- 
ulus deltoides (211). [foliose/corticolous]. W. 

Physcia millegrana Degel. Common; on the trunks and lower branches of 
Populus deltoides, Quercus velutina (198), and Salix interior (213). 


Physcia stellaris (L.) Nyl. Frequent; on the trunks and lower branches of 
Populus deltoides and Quercus velutina (209). [foliose/corticolous]. 

Punctelia Krog 

Punctelia rudecta (Ach.) Krog. Rare; on the trunk of Quercus rubra (205). 


Thelidium Mas sal. 

Thelidium microcarpum (Leight.) A.L. Sm. Rare; on concrete rubble 


Xanthoria (Fr.) Th. Fr. 

Xanthoma fallax (Hepp) Arnold. Occasional; on the lower branches of Quer- 
cus velutina (814) and on a wood rail fence (251). [foliose / corticolous 


Proceedings of the Indiana Academy of Science 3 3 

(1997) Volume 106 p. 33-37 





David Polley 

Biology Department 

Wabash College 

Crawfordsville, Indiana 47933 

ABSTRACT: The genetic evidence for two independent potassium transport 
systems in Chlamydomonas reinhardtii is presented. The first system was pre- 
viously described and is encoded by the TRK genes. The second, described 
here, is encoded by the HKR gene. 

KEYWORDS: Chlamydomonas reinhardtii, potassium transport, transport 


Potassium is the major monovalent cation in plant and algal cells, where it 
plays an important role in several cellular processes, such as osmoregulation, 
protein synthesis, and charge balance (Leigh and Wyn- Jones, 1984). Because of 
the essential role potassium plays in these processes, its concentration within the 
cytoplasm and various other cell compartments is highly regulated. This control 
is achieved through the regulation of potassium transport across the membrane 
barriers of various compartments. For example, the cytoplasmic potassium con- 
centration is maintained at a fairly constant level of approximately 150 mM even 
though plants grow successfully in media with potassium concentrations rang- 
ing from 10 jitM to 10 mM. A higher plant's ability to respond to a range of exter- 
nal potassium concentrations is achieved by two kinetically distinct transport 
systems. The first, a high-affinity system, exhibits typical Michaelis-Menten 
kinetics and reaches saturation in the micromolar range; the second, a low-affin- 
ity system, operates in the millimolar range and is often said to be non-saturable 
(Kochian and Lucas, 1982). In a recent study of potassium transport in the green 
alga Chlamydomonas reinhardtii, Malhotra and Glass (1995) reported kinetic 
data that support the existence of two potassium transport systems, a saturable, 
high-affinity system (HATS) and a non-saturable, low-affinity system (LATS). 

In recent years, much progress in our understanding of the nature of potas- 
sium transporters has been made through a variety of kinetic and electrophysi- 
ological analyses (e.g., Kochian and Lucas, 1982; Maathius and Sanders. 1994). 
Insight into the structure of transporters has been enhanced greatly with the 
cloning of plant potassium channel genes (Anderson, et ah, 1992; Sentenac. 
et al, 1992). This progress notwithstanding, gaps remain in our understanding. 

34 Cell Biology: Polley Vol. 106 (1997) 

One approach to the study of potassium transport that has not been explored 
fully is the isolation and characterization of potassium transport defective clones. 
This approach, employing Chlamydomonas as a model system, was first out- 
lined by Polley and Doctor (1985), who isolated and conducted a preliminary 
characterization of three potassium transport defective clones. Subsequently, 
additional mutant clones have been isolated and characterized, and the presence 
of at least three unlinked genes (trk) that encode the high affinity transport sys- 
tem has been demonstrated (Polley, in review). In work reported here, genetic 
evidence for a second potassium transport system is presented. Isolation of mutant 
transport alleles should make possible the identification of additional compo- 
nents of the potassium transport system and the future cloning of genes involved 
in transport and its regulation. 


Strains. The mutant clone DP2 (trk2-l actl mt-) was derived as a recom- 
binant from a cross between the potassium transport defective clone KDP5 (trk2- 
1 mt+) and CC1680 (actl ac80 mt-). KDP5 was isolated after UV-mutagenesis 
of wild-type strain 137c (Polley, in review). Both the wild-type and mutant strain 
CC1680 were obtained from the Chlamydomonas Genetics Center. Mutant clone 
10KDP1 was isolated after UV-mutagenesis of clone DP2. 

Media and Growth Conditions. Cells were grown axenically at 25° C under 
continuous illumination either in aerated liquid cultures or on medium solidified 
with agar. Medium 0K0N is a modification of tris-acetate-phosphate medium 
(TAP medium) that possesses only trace amounts of potassium or sodium (Pol- 
ley and Doctor, 1985). The potassium requirement of mutant clones was deter- 
mined by measuring the rate of growth in 0K0N supplemented with different 
concentrations of KC1. Growth was monitored by following absorbance (light 
scattering) at 560 nm. Absorbance is directly proportional to cell number, and 
the same correlation between cell number and absorbance exists for cells grown 
in media of different potassium concentrations. 

Mutagenesis and Genetic Analysis. Four milliliters of DP2 cells grown 
to a density of approximately 5 x 10 6 cells/ml were placed in a 4.5 cm Falcon 
petri dish and exposed to UV-irradiation for 90 sec in a Strategene UV Crosslink- 
er. After exposure, cells were grown overnight in the dark to fix the mutation. 
The cells were then plated on 10K0N (0K0N medium supplemented with 10 mM 
KC1). Approximately 5-10% of the cells survive UV irradiation. Survivors were 
screened by replica-plating to IKON (0K0N supplemented with 1 mM KC1). 
Mating and tetrad analysis were done as described by Harris (1989). 


Approximately 1 ,000 DP2 clones that survived UV-mutagenesis were screened 
for their ability to grow on IKON. Of these, 5 failed to grow, and they were 
designated as putative high K + requiring (hkr) mutants. One of these, clone 
10KDP1 , was selected for further study. The concentration of potassium required 

Vol. 106 (1997) Indiana Academy of Science 35 

Table 1. Growth rate of wild-type and mutant clones (growth rate constants, k, where 
At = A e kt , are the averages of three experiments ± standard error). 

Growth Rate Constant (k x 10 2 ) 












trk2-l HKR 
trk2-l hkr 









* ND = not done. 

for growth and the specificity of the requirement were determined by measur- 
ing the rate of growth in liquid media supplemented with different concentra- 
tions of KC1 (Table 1). Growth was monitored by measuring absorbance (see 
methods); readings were taken every 3-5 hours during exponential growth of the 
culture. The growth rate constant, k, is based on 6 time points. The reported val- 
ues in Table 1 are the averages of three experiments. As the data clearly show, 
trk2-l cells require KC1 concentrations greater than 0.1 mM to achieve a wild- 
type growth rate; trk2-l hkr cells require concentrations greater than 1 mM. In 
both cases, NaCl cannot substitute for KC1 (Polley and Doctor, 1985, and data 
not shown). 

The trk2-l allele maps to linkage group II and is linked to the actl allele 
(PD > NPD, Table 2, Tetrad Analysis; Polley, in review). The mutant clone 
lOKDPl was crossed to wild type (trk2-l hkr actl mt- x TRK2-1 HKR ACT1 
mt+), and tetrad analysis was performed in order to map the genetic lesion, hkr, 
relative to the trk2-l and actl alleles. The possible genotypes resulting from this 
cross and their respective phenotypes and tetrads are shown in Table 2 (Geno- 
types and Phenotypes). The phenotypes are defined as low KC1 when cells achieve 
wild-type growth rate on 0. 1 mM KC1, as intermediate KC1 when cells require 
1 mM KC1, and as high KC1 when cells require 10 mM KC1 in order to grow at 
wild-type rates. Of the four genotypes listed, only the phenotype of the recom- 
binant TRK2-1 Mr could not be predicted a priori. The wild-type TRK2-1 allele 
was assumed to be epistatic to hkr, and, therefore, the phenotype of TRK2-1 
hkr would be low KC1. That this assumption is correct is supported by the obser- 
vation that only three classes of tetrads, as defined by the phenotypic ratios of 
tetrad products, were obtained (Table 2, Tetrads and Phenotypes). Based on the 
criteria PD = NPD and NPD/T > 0.25 (Perkins, 1953), the tetrad data (Table 2, 
Tetrad Analysis) show that the hkr gene is unlinked to trk.2-1. 

While the hkr lesion in lOKDPl might actually be a mutation in one of the 
other two, unlinked TRK genes, this possibility seems unlikely for two reasons. 
First, genetic recombinants harboring two mutant TRK alleles (trkl trk2-l, trkl 
trk3, or trk2-l trk3) do not exhibit growth rates different from cells with just one 
mutant TRK allele (Polley, in review). This finding suggests that the TRK gene 


Cell Biology: Polley 

Vol. 106 (1997) 

Table 2. Genetic analysis of clone 10KDP1 

Genotypes and Phenotypes 

Genotype Phenotype 


trk2-l hkr 

trk2-l HKR 

TRK2-1 hkr 

Low KC1 

High KC1 

Intermediate KC1 

Low KC1 

Tetrads and Phenotypes 

Tetrads Phenotypic Ratios 

Parental Ditype 2 High : 2 Low 

Nonparental Ditype 2 Inter. : 2 Low 

Tetratype 2 Inter. : 1 Low : 1 High 

Tetrad Analysis 


trk2-l actl trk2-l hkr 

products interact functionally; 
i.e., if one mutant polypeptide 
disrupts function of a multimeric 
protein complex, a second mutant 
polypeptide would not have a 
noticeable effect because the 
complex is already disabled. The 
hkr allele in a trk2-l background, 
however, does increase the potas- 
sium requirement for growth. 
The hkr allele must, therefore, 
affect the function or regulation 
of some other potassium trans- 
port system. 

Second, and consistent with 
the above interpretation, is the 
observation that TRK2-1 is epista- 
tic to hkr. If the TRK genes encode 
for a potassium transport sys- 
tem that is able to transport potas- 
sium from media with low 
concentrations (0.1 mM) of potas- 
sium, and if the HKR gene encodes for a transport system that is effective only 
in media with intermediate levels (1.0 mM) of potassium, then one would expect 
trk hkr cells to require high concentrations (10 mM) of potassium to grow. By 
similar reasoning, one would also expect cells with a functional TRK system not 
to need an HKR system; i.e., TRK2-1 hkr cells will grow on low levels of potas- 

In summary, the above genetic data support the conclusion that two distinct 
transport systems exist: the TRK system which operates at low levels of potas- 
sium and the HKR system which operates at higher levels of potassium. Future 
work will involve a kinetic analysis of potassium transport by 10KDP1, map- 
ping the hkr mutant allele, and determining if hkr in a trkl or trk3 background 
also require, as would be expected, high concentrations of potassium. 

10KDP1 xCC125 

16:0: 5 



Anderson, J. A., S.S. Huprikar, L.V. Kochian, W.J. Lucas, and R.F. Gaber. 1992. Functional expression of a 
probable Arabidopsis thaliana potassium channel in S. cerevisiae. Proc. Nat. Acad. Sci. USA 49: 684- 

Harris, E.H. 1989. The Chlamydomonas source book. Academic Press, Inc., San Diego, California, 780 pp. 

Kochian, L.V. and W.J. Lucas. 1982. Potassium transport in corn roots. 1. Resolution of kinetics into a sat- 
urable and linear components. Plant Physiol. 70: 1723-1731. 

Leigh, R.A. and R.G. Wyn-Jones. 1984. A hypothesis relating critical potassium concentrations for growth 
to the distribution and functions of this ion in the plant cell. New Phytol. 97: 1-13. 

Maathius, F.J.M. and D. Sanders. 1994. Mechanisms of high-affinity potassium uptake in roots of Arabidop- 
sis thaliana. Proc. Nat. Acad. Sci. USA 91: 9272-9276. 

Vol. 106 (1997) Indiana Academy of Science 37 

Malhotra, B. and A.D.M. Glass. 1995. Potassium fluxes in Chlamydomonas reinhardtii. 1 . Kinetics and elec- 
trical potentials. Plant Physiol. 108: 1527-1536. 

Perkins, D.D. 1953. The detection of linkage in tetrad analysis. Genetics 38: 187-197. 

Polley, L.D. and D.D. Doctor. 1985. Potassium transport in Chlamydomonas reinhardtii: Isolation and char- 
acterization of transport deficient mutant strains. Planta 163: 208-213. 

Sentenac, H., N. Bonneaud, M. Minet, F. Lacroute, J.-M. Salmon, F. Gaymard, and C. Grignon. 1992. Cloning 
and expression in yeast of a plant potassium ion transport system. Science 256: 663-665. 

Proceedings of the Indiana Academy of Science 39 

(1997) Volume 106 p. 39-51 





Shannon E. Brown 

School of Forestry and Wood Products 

Michigan Technological University 

Houghton, Michigan 4993 1 


George R. Parker 

Department of Forestry and Wood Products 

Purdue University 

West Lafayette, Indiana 47906 

ABSTRACT: The white-tailed deer (Odocoileus virginianus) is an herbivo- 
rous species that can significantly influence the structure of forest communi- 
ties wherever it occurs in excessive numbers. This species has a propensity to 
increase beyond the carrying capacity of its habitat when predation pressure 
is reduced. The influence of white-tailed deer on the plant communities with- 
in Indiana's State Parks, which have been closed to hunting by man for sev- 
eral decades, are explored in this paper. During 1993 and 1994, sites (stratified 
by physiographic position) were sampled within Brown County State Park 
and adjacent State and National Forests (open to public hunting). A reduction 
in the percentage cover of the ground flora and the mature height of some spe- 
cific plant species inside Brown County State Park as compared to external 
sites subjected to hunting pressure was noted. In addition, the recruitment of 
many woody species into larger size classes was reduced significantly with- 
in Brown County State Park. The current forest structure within Brown Coun- 
ty State Park indicates that this reduction in recruitment has been occurring 
for many years. The reduction in recruitment, along with the damage docu- 
mented within the herbaceous layer, suggests that the sustainability and struc- 
ture of the forests within the park may ultimately be affected unless deer 
numbers are controlled and the forest understory is allowed to recover. 

KEYWORDS: Adiantum pedatum, Brown County State Park, Indiana, per- 
centage cover, plant communities, species richness, white-tailed deer. 


White-tailed deer, along with large predators, had been eliminated from Indi- 
ana by 1900 (Mumford and Whitaker, 1982). In 1934, the Federal Aid to Wildlife 
Restoration (Pittman-Robertson Act) provided funds for the reintroduction of 
white-tailed deer into southern Indiana (Mumford and Whitaker, 1982). These 
introduced deer benefitted from a landscape of second-growth forests, openings, 
and farmlands that had been created by logging, clearing, and agriculture (Smith. 

40 Ecology: Brown and Parker Vol. 106 (1997) 

By 1948, the Pittman-Robertson project reported an overabundance of 
deer and deer-induced damage to agricultural and native plant communities, lead- 
ing to the first modern hunt within 17 Indiana counties in 1951 (Mumford and 
Whitaker, 1982). State Parks in Indiana were closed to hunting and have remained 
so until recently. Restricted hunting has allowed the white-tailed deer popula- 
tions in some State Parks to expand beyond the capacity of their habitat to sup- 
port them. 

Deer have a selective foraging strategy when food resources are abundant, 
but they change to a more generalist strategy as resources are depleted (Brown 
and Doucet, 1991; Strole and Anderson, 1992; Kohlman and Risenhoover, 1994). 
Selective foraging by a generalist species is a learned behavior which maximizes 
nutrient intake but is subject to constraints at low levels of food availability 
(Westoby, 1974). Deer are known to select a species according to its chemical 
composition, and intake has been found to be constrained by forage toxicity 
(Mc Arthur, et ai, 1993). These foraging strategies often lead to browse species 
preference. Several studies show that deer select certain herbaceous and 
woody species over others (Korschgen, 1962; Nixon, et ai, 1970; McCaffery, 
et ai, 1974). This foraging strategy is termed the "optimization model," and the 
model implies that the grazing pressure on a species within a plant community 
may increase as that species becomes rarer (Westoby, 1974). 

Foraging strategies have been found to be sensitive to changes in plant avail- 
ability (Kohlman and Risenhoover, 1994). Deer avoid nonpreferred browse 
species even when more preferred species become depleted. Deer search for and 
consume preferred forage until it becomes rare before switching to an apparently 
less desired species (Gillingham and Bunnell, 1989). Deer density may also affect 
food selection. Preferred foods, such as acorns, were found to be utilized more 
quickly by a larger deer population (McCullough, 1985; Kohlman and Risen- 
hoover, 1994). Therefore, at higher population levels, deer could significantly 
alter plant communities by extirpating the more preferred species. 

Evidence of intensive browsing by white-tailed deer in forest plant com- 
munities has been documented (Marquis, 1974; Alverson, et al., 1988; Strole 
and Anderson, 1992; Balgooyen and Waller, 1995) and found to negatively effect 
the regeneration of some tree species by reducing their recruitment and survival 
(Marquis, 1974; Strole and Anderson, 1992). This effect on regeneration has 
been studied in the eastern hemlock (Tsuga canadensis) forests of northeastern 
Wisconsin, where, in areas that are heavily browsed, sugar maple (Acer sac- 
charum) rapidly replaces hemlock (Anderson and Loucks, 1979). However, 
Anderson and Loucks (1979) also found that once browsing pressure was relieved, 
the hemlock recovered. Their finding suggests that a species subject to intensive 
browsing may recover if the pressure is reduced. In addition, the amount of 
browsing pressure on certain preferred forage species could be used as an indi- 
cator of deer abundance within an area (Korschgen, 1962). 

In 1993, a committee was established by the Indiana Department of Natur- 
al Resources to develop alternative solutions for managing the excessive popu- 

Vol. 1 06 ( 1 997) Indiana Academy of Science 4 1 

lation of white-tailed deer within Brown County State Park, Indiana. The com- 
mittee concluded that the deer were seriously damaging understory vegetation 
and recommended a limited hunt within the park boundaries. This hunt occurred 
in December 1993. The committee also initiated a study to investigate the effects 
of overbrowsing on understory vegetation (Brown County State Park Deer Study 
Committee, 1993). The forest communities within Brown County State Park, 
Yellowwood State Forest, and Hoosier National Forest (the latter two are both 
open to hunting) were sampled in 1993 by James Van Kley (a Ph.D. student at 
Purdue University) to determine the impact of white-tailed deer on the vegeta- 
tion. Preliminary measurements of the structure of the plant community and 
the browse damage within Brown County State Park indicated that the deer were 
browsing heavily on selected herbaceous and woody species and significantly 
reducing the percentage cover, species richness, and mature heights of the ground 
flora species. This study also found a difference between the southwestern cor- 
ner and the main body of Brown County State Park, with the southwestern cor- 
ner showing less damage (Parker and Van Kley, 1993). Parker and Van Kley 
(1993) theorized that because the southwestern corner was bordered by Yel- 
lowwood State Forest and Hoosier National Forest, the deer in the southwestern 
corner of Brown County State Park experienced more hunting pressure than those 
within the main body of the park. 

Permanent study areas were not established by Van Kley to observe trends 
in browse damage. Therefore, the current study was initiated in the spring of 
1994 to determine the structure and composition of the plant communities fol- 
lowing the removal of 392 deer in December 1993. Permanent plots were estab- 
lished to monitor changes in the plant communities in response to long-term 
change in deer populations. 


Brown County State Park, located in south-central Indiana, was first opened 
in 1929 and is Indiana's largest State Park at 6,358 hectares. The park is locat- 
ed within the Brown County Hills Section of the Highland Rim Natural Region 
(Homoya, et al., 1985). This area is defined by acid silt loam soils with a small 
amount of loess and characteristic topographic features such as deeply dissect- 
ed uplands, steep slopes, and narrow hollows (Homoya, et aL, 1985; Van Kley 
and Parker, 1993). Uplands are dominated by oak-hickory forest (Homoya. et 
al, 1985); common plants on the upper slopes are chestnut oak (Quercus pri- 
nus), common greenbrier (Smilax rotundifolia), low growing shrubs, and sedges 
(in particular, painted sedge {Carex picta)). The ravines contain mesic species 
(Homoya, et ah, 1985) such as beech (Fagus grandifolia), red oak {Quercus 
rubra), sugar maple (Acer saccharum), and white ash (Fraxinus americana). 
Control areas were selected within Yellowwood State Forest and the Pleasant 
Run Unit of the Hoosier National Forest, which border the southwestern section 
of Brown County State Park. Both the State and National Forests are open to 
annual deer harvest. 

42 Ecology: Brown and Parker Vol. 106 (1997) 


Twenty sites within Brown County State Park and ten control sites in Yel- 
lowwood State Forest/Hoosier National Forest were randomly selected to estab- 
lish permanent study plots. The 20 sites in the park were divided into eight 
sites in the southwestern corner, where less browsing was observed (Parker 
and Van Kley, 1993), and 12 in the main body of the park. The Ecological 
Classification System developed for the Pleasant Run Unit of Hoosier Nation- 
al Forest was used to determine similar physiographical units inside and outside 
the park (Van Kley and Parker, 1993). Sites were limited to closed-canopy, rel- 
atively mature forests and divided into four ecological landtypes: mesic norther- 
ly or northeasterly slopes, dry southerly or southwesterly slopes, mesic bottomlands, 
and dry ridges. One or a combination of these landtypes were sampled at each 
site; a total of 19 landtypes were sampled in Yellowwood State Forest/Hoosier 
National Forest, 16 within the southwestern corner of Brown County State Park, 
and 27 in the main body of Brown County State Park. All the sites were sam- 
pled within two months of each other to avoid site-to-site variation. 

The following vegetative variables were measured using a series of nested 
plots at each sample location: the percentage cover of the ground flora, the species 
richness of the ground flora, and the stem density and species richness of the 
woody species within four size classes (1 = < 50 cm in height; 2 = 50 to 200 
cm in height; 3 = > 200 cm in height and < 2.5 cm dbh; and 4 = > 200 cm in 
height and > 2.5 cm dbh). The mature heights of six herbaceous species were 
also recorded. 

The percentage cover of the ground flora was measured using a ten-meter 
line transect running parallel to the contour of the slope in each landtype mea- 
sured at a site. The length of overlap (cm) for the ground flora, including both 
herbaceous and woody class 1 , was recorded and used to calculate the percent- 
age cover for each species. 

To record the number of stems and woody species in classes 1 and 2, a one 
meter by ten meter belt transect was established using the ten-meter line tran- 
sect as its lower edge. The dbh of woody species in classes 3 and 4 was record- 
ed within a 100 m 2 circular plot, the center of which was the start of the line 

Ground flora richness in each landtype was determined by recording all 
the herbaceous species found within the belt transect and circular plot. The species 
present within the circular plot were determined by walking concentric circles 
from the outer edge to the center. 

Six species were chosen to test for differences in mature plant height with- 
in Yellowwood State Forest/Hoosier National Forest, the southwestern corner 
of Brown County State Park, and the main body of Brown County State Park. 
The chosen species showed dramatic differences in mean mature height in heav- 
ily browsed areas relative to less impacted ones (Parker and Van Kley, 1993). 
The species measured were: maidenhair fern (Adiantum pedatum), Jack-in-the- 
pulpit (Arisaema triphyllum), wild licorice {Galium circaezans), sweet cicely 

Vol. 1 06 ( 1 997) Indiana Academy of Science 43 

(Osmorhiza claytonii), Christmas fern {Polystichum acrostichoides), and com- 
mon greenbrier (Smilax rotundifolia). To determine mature height, the forbs were 
measured along the stem to the node with the highest leaf or whorl of leaves. For 
Christmas fern, the length of the longest frond was measured. Three to five plants 
of each species were measured within each belt transect and their heights were 
averaged to determine the mean height of the species. The percentage cover was 
also calculated for these species. 

One-way analysis of variance was used to compare the means for vegeta- 
tion variables among the sites within Yellowwood State Forest/Hoosier 
National Forest, the southwestern corner of Brown County State Park, and the 
main body of Brown County State Park. For tests of mature height and percentage 
cover of the individual species, only the landtypes in which the species were 
known to be common were used. These included mesic slopes and bottom- 
lands for maidenhair fern, Jack-in-the-pulpit, wild licorice, sweet cicely, and 
Christmas fern. All sites were used for common greenbrier. An alpha level of 
P < 0. 10 was used to delineate significant differences in the vegetation variables 
among the three sample areas. 


The average number of species was not significantly different in the three 
areas (Table 1). However, on average, more species were found in all landtypes 
in Yellowwood State Forest/Hoosier National Forest and the southwestern cor- 
ner of Brown County State Park than in the main body of the park. The per- 
centage cover of the ground flora was significantly reduced in the main body 
of the park compared to both its southwestern corner and the areas outside the 
park. The coverage of the ground flora averaged only 14.4% per sample site in 
all landtypes within the main body of Brown County State Park as compared to 
27.8% in the park's southwestern corner and 21.7% in Yellowwood State For- 
est/Hoosier National Forest. The greatest difference in percentage cover of the 
ground flora was found on mesic and bottomland sites. Coverage within the main 
body of Brown County State Park was 50% less than in Yellowwood State For- 
est/Hoosier National Forest. 

Specific species exhibited differences in their percentage cover within the 
park when compared to the control areas (Table 2). Maidenhair fern and sweet 
cicely both exhibited a significantly lower percentage cover inside the south- 
western corner and in the main body of the park when compared to Yellowwood 
State Forest/Hoosier National Forest. Sweet cicely did not intersect the tran- 
sect within the park's southwestern corner at any sites and only overlapped the 
transect at 0.3% of the sites in the main body of the park as compared to 1.5% 
in Yellowwood State Forest/Hoosier National Forest. Christmas fern showed a 
significantly higher percentage cover in Yellowwood State Forest/Hoosier Nation- 
al Forest and the southwestern corner of the Brown County State Park relative 
to the main body of the park, where it averaged 0.9% per sample site. Common 
greenbrier had a significantly higher percentage cover in Yellowwood State For- 


Ecology: Brown and Parker 

Vol. 106 (1997) 

Table 1. Mean ground flora richness measured within a 10 m 2 belt transect and a 100 
m 2 circular plot, and the percentage cover of the ground flora measured on a 10 m line 
transect observed at sample locations outside Brown County State Park, in the south- 
western corner of Brown County State Park, and in the main body of Brown County 
State Park during 1994. Row means with different letters are significantly different 

Variable Sampled at Each Yellowwood State and 
Ecological Landtype Hoosier National Forests 

Southwestern Corner of 
Brown County State Park 

Main Body of Brown 
County State Park 

All Sites (n = 62) 

Ground flora richness 
Percentage cover of ground flora 

21.5 a 

21.7 a 

27.8 a 

19.8 a 
14.4 b 

Mesic Bottomlands (n = 15) 

Ground flora richness 
Percentage cover of ground flora 

32.0 a 

33.7 a 

30.1 a 
40.9 a 

28.9 a 
14.5 b 

Mesic Slopes (n = 22) 

Ground flora richness 
Percentage cover of ground flora 

24.6 a 
18.0 a 

24.2 a 
22.0 a 

21.5 a 
8.0 b 

Dry Slopes (n = 15) 

Ground flora richness 
Percentage cover of ground flora 

13.6 a 
22.3 a 

14.0 a 
34.9 a 

12.8 a 

Dry Ridges (n = 10) 

Ground flora richness 
Percentage cover of ground flora 

7.5 a 

15.3 a 
14.7 a 

13.0 a 
12.2 a 

est/Hoosier National Forest than in either area of the park. The percentage cover 
of greenbrier was significantly greater in the southwestern corner relative to 
the main body of Brown County State Park. Jack-in-the-pulpit and wild licorice 
displayed no significant differences in coverage among the three areas. 

The mean heights of the mature plants for the six selected species were 
also compared (Table 2). Jack-in-the-pulpit and common greenbrier were found 
to be significantly taller in Yellowwood State Forest/Hoosier National Forest 
than in both the southwestern corner and main body of Brown County State Park. 
Maidenhair fern and Christmas fern were significantly shorter in the main 
body of Brown County State Park when compared to the southwestern corner 
of the park and Yellowwood State Forest/Hoosier National Forest. Both Jack- 
in-the-pulpit and maidenhair fern were approximately 10 cm taller within Yel- 
lowwood State Forest/Hoosier National Forest than in both the southwestern 
corner and the main body of the park. Common greenbrier averaged 45 cm taller 
in Yellowwood State Forest/Hoosier National Forest when compared to both 

Vol. 1 06 ( 1 997) Indiana Academy of Science 


Table 2. The mean percentage cover and mean height for selected species observed with- 
in a 10 m 2 belt transect at sample locations outside Brown County State Park, in the 
southwestern corner of Brown County State Park, and in the main body of Brown Coun- 
ty State Park during 1994. The values used are from landtypes where the species was 
common. Row means for heights and percentage cover of the ground flora with differ- 
ent letters are significantly different (P < 0.10). 

Critical Species and Yellowwood State and 
Variable Tested Hoosier National Forests 

Southwestern Corner of 
Brown County State Park 

Main Body of Brown 
County State Park 

Adiantum pedatum ' 

Percentage cover 
Height (cm) 

2.3 a 
31.1 a 

0.4 b 
32.2 a 

22.0 b 

Arisaema triphyllum 1 

Percentage cover 
Height (cm) 

0.9 a 

22.3 a 

0.9 a 
12.9 b 

1.2 a 
12.0 b 

Galium circaezans 1 

Percentage cover 
Height (cm) 

15.4 a 

0.07 a 
15.0 a 

0.08 a 
14.6 a 

Osmorhiza claytoniV 

Percentage cover 
Height (cm) 

1.5 a 

35.2 a 

24.0 ab 

0.3 b 

Polystichum acrostichoides x 

Percentage cover 
Height (cm) 

4.7 a 
51.5 a 

2.8 a 
41.9 a 

0.9 b 
26.5 b 

Smilax rotundifolia 2 

Percentage cover 
Height (cm) 

4.9 a 

95.9 b 


1 Mesic slopes and bottomlands. 

2 All sites. 

areas within Brown County State Park. Sweet cicely was significantly taller in 
Yellowwood State Forest/Hoosier National Forest than in the main body of the 
park, averaging 35.2 cm within Yellowwood State Forest/Hoosier National For- 
est and 11.2 cm within the main body of Brown County State Park. Wild licorice 
displayed no significant difference in mean height among the three sampling 

Overall, the number for woody stems (class 1) was significantly greater in 
the southwestern corner of Brown County State Park relative to Yellowwood 
State Forest/Hoosier National Forest and the main body of the park (Table 3). 
The greatest difference occurred on mesic slopes and dry ridges where the num- 

46 Ecology: Brown and Parker Vol. 106 (1997) 

Table 3. The number of stems for woody species and the species richness within four 
size classes recorded at sample sites outside of Brown County State Park, in the south- 
western corner of Brown County State Park, and in the main body of Brown County 
State Park during 1994. Row means with different letters are significantly different 
(P < 0.10). Size class 1 = stems < 50 cm in height, size class 2 = stems 50 to 200 cm in 
height, size class 3 = stems > 200 cm in height and < 2.5 cm dbh, and size class 
4 = stems > 200 cm in height and > 2.5 cm dbh. The density of size classes 1 and 2 are 
per 10 m 2 plot and for size classes 3 and 4 are per hectare. 

Critical Species and 

Yellowwood State and 

Southwestern Corner of 

Main Body of Brown 

Variable Tested 

Hoosier National Forests 

Brown County State Park 

County State Park 

All Sites 

Size class 1 


39.6 a 

67.6 b 

39.4 a 

Species richness 

8.4 a 

8.8 a 


Size class 2 


1.3 a 

0.6 ab 

0.2 b 

Species richness 

0.8 a 

0.5 a 

0.04 b 

Size class 3 


405 a 

94 b 

159 b 

Species richness 

1.8 a 

0.7 b 

1.0 b 

Size class 4 


932 a 


833 a 

Mesic Bottomlands 

Size class 1 


47.0 a 

45.3 a 


Species richness 

8.5 a 

10.0 a 

5.9 b 

Size class 2 


1.0 a 

0.3 a 

0.6 a 

Species richness 

1.0 a 

0.2 b 

0.14 b 

Size class 3 


200 a 

75 a 

143 a 

Species richness 

1.0 a 

0.8 a 

0.9 a 

Size class 4 


700 a 

750 a 


Mesic Slopes 

Size class 1 


38.9 a 

90.0 b 

30.3 a 

Species richness 

8.1 a 

7.3 a 

5.5 a 

Size class 2 


2.0 a 

0.8 ab 


Species richness 

1.0 a 

0.6 a 


Size class 3 


450 a 

150 b 

175 b 

Species richness 

2.4 a 

0.8 b 


Size class 4 


875 a 

833 a 

838 a 

Vol. 106 (1997) Indiana Academy of Science 


Table 3 (Continued) 

Critical Species and 

Yellowwood State and 

Southwestern Corner of 

Main Body of Brown 

Variable Tested 

Hoosier National Forests 

Brown County State Park 

County State Park 

Dry Ridges 

Size class 1 


18.0 a 

73.3 b 

25.0 a 

Species richness 

6.0 a 

9.0 b 

5.6 a 

Dry Slopes 

Size class 1 


43.4 a 

46.7 a 


Species richness 

9.8 a 

9.7 a 

7.3 a 

Dry Slopes and Ridges 

Size class 2 


0.7 a 

0.7 a 


Species richness 

0.6 a 

0.5 a 


Size class 3 



50 b 

142 c 

Species richness 

1.7 a 

0.5 b 

0.9 b 

Size class 4 


1128 a 

883 a 

925 a 

ber of stems in the southwestern corner of the park averaged 90.0 and 73.3, 
respectively. Dry slopes and mesic bottomlands did not exhibit any significant 
differences among the three areas. 

The number of species observed in the one meter by ten meter belt transect 
was significantly higher for all landtypes in Yellowwood State Forest/Hoosier 
National Forest and the southwestern corner of Brown County State Park than 
in the main body of the park. The difference was most evident in the mesic bot- 
tomland sites in Brown County State Park which averaged 5.9 species per sam- 
ple site compared to 8.5 in Yellowwood State Forest/Hoosier National Forest 
and 10.0 in the southwestern corner of the park. The other landtypes did not show 
any significant differences with the exception of dry ridges, where a significantly 
higher number of species was observed in the southwestern corner of Brown 
County State Park than in either Yellowwood State Forest/Hoosier National For- 
est or the main body of the park. 

The overall number of stems for all woody species (class 2) per sample site 
for all landtypes was significantly higher in Yellowwood State Forest/Hoosier 
National Forest when compared to the main body of the park (Table 3). Sites 
averaged 1,300 stems per hectare in Yellowwood State Forest/Hoosier Nation- 
al Forest as compared to 200 stems per hectare inside the main body of the park. 
Mesic slopes as well as dry slopes and ridges also exhibited this pattern: how- 
ever, significant differences were not observed for these landtypes in the south- 
western corner of Brown County State Park relative to either Yellowwood 

48 Ecology: Brown and Parker Vol. 106 (1997) 

State Forest/Hoosier National Forest or the main body of the park. Mesic bot- 
tomlands did not show any significant differences among the three areas. The 
number of woody species in size class 2 was also significantly higher in Yel- 
lowwood State Forest/Hoosier National Forest and the southwestern corner of 
Brown County State Park than in the main body of the park, where papaw (Asim- 
ina triloba) was the only species found on transects (Table 3). 

The number of woody stems (class 3) per hectare was significantly reduced 
on all sites except mesic bottomlands within the park when compared to Yel- 
lowwood State Forest/Hoosier National Forest (Table 3). Species richness for 
this size class followed the same pattern (Table 3). Significantly more species 
per sample site were found within Yellowwood State Forest/Hoosier National 
Forest, which averaged 1.8 species per sample site, than in the southwestern cor- 
ner (0.7) or the main body (1.0) of the park. 

The number of woody species (class 4) per hectare was not significantly dif- 
ferent inside Brown County State Park when compared to Yellowwood State 
Forest/Hoosier National Forest (Table 3). The number of stems averaged 932 
per hectare for all landtypes within Yellowwood State Forest/Hoosier National 
Forest as compared to 831 in the southwestern corner and 833 in the main 
body of the park, respectively. This pattern was observed for all the different 

The same vegetation variables were also measured in 1993 within Brown 
County State Park and Yellowwood State Forest/Hoosier National Forest by Van 
Kley, and the means of these variables show similar trends between the two years 
(Table 4). For example, the average number of ground flora species per site found 
outside the park in 1994 was similar to the number found in 1993 (21.5 in 1994 
and 23.3 in 1993). However, the number of species found per site in the south- 
western corner and main body of Brown County State Park was higher in 
1994. Percentage cover of the ground flora was similar in both years within the 
main body of Brown County State Park, averaging 14.4% in 1994 and 14.7% 
in 1993. However, Yellowwood State Forest/Hoosier National Forest exhibited 
a slightly lower percentage cover, and the southwestern corner of Brown 
County State Park exhibited a higher percentage cover in 1994. The southwest- 
ern corner of Brown County State Park also exhibited an increase in the num- 
ber of seedlings in 1994. In contrast, sapling numbers stayed constant between 
the two sampling seasons. 


Excessive browsing within Brown County State Park has resulted in reduced 
numbers and sizes for some plant species, a conclusion consistent with those of 
Alverson, et al. (1988) and Balgooyen and Waller (1995). These researchers 
found that white-tailed deer could directly and indirectly affect the abundance 
and structure of many herbaceous species, thereby influencing forest composi- 
tion. Likewise, Balgooyen and Waller (1995) indicated that many herbaceous 
species, especially preferred browse species, were jeopardized by high deer den- 

Vol. 106 (1997) Indiana Academy of Science 


Table 4. The mean species richness and percentage cover of the ground flora as well as 
the mean density of woody stems in size classes 1 and 2 recorded at sample sites out- 
side Brown County State Park, in the southwestern corner of Brown County State 
Park, and in the main body of Brown County State Park during 1993 (Parker and Van 
Kley, 1993). Row means with different letters are significantly different (P < 0.10). Size 
class 1 = stems < 50 cm in height, and size class 2 = stems 50 to 200 cm in height. 

Critical Species and Yellowwood State and Southwestern Corner of Main Body of Brown 
Variable Tested Hoosier National Forests Brown County State Park County State Park 

All Sites 

Ground flora richness 
Percentage cover of ground flora 
Woody size class 1 
Woody size class 2 

Mesic Bottomlands 

Ground flora richness 
Percentage cover of ground flora 
Woody size class 1 
Woody size class 2 

Mesic Slopes 

Ground flora richness 
Percentage cover of ground flora 
Woody size class 1 
Woody size class 2 

Dry Slopes 

Ground flora richness 
Percentage cover of ground flora 
Woody size class 1 
Woody size class 2 

23.3 a 

16.6 b 

14.7 b 

34.1 a 

18.6 b 

14.6 b 

33.3 a 

23.7 b 

13.9 c 

3.9 a 

0.5 b 

0.4 b 

34.8 a 

22.4 b 

20.8 b 

39.0 a 

28.3 a 


47.2 a 

13.2 b 

8.8 b 

5.0 a 

0.6 b 

0.5 b 

30.3 a 

21.3 b 


32.4 a 

10.9 b 

8.0 b 

35.8 a 

16.9 b 


4.0 a 


0.5 b 

14.0 a 

12.1 ab 

9.9 b 

29.2 a 

23.6 a 

21.2 a 

25.6 ab 

33.9 a 

16.8 b 

4.6 a 

0.8 b 

0.2 b 

sities. The significant reduction in both percentage ground cover and the size 
of several herbaceous species within Brown County State Park when com- 
pared to Yellowwood State Forest/Hoosier National Forest indicates that the deer 
population had exceeded the carrying capacity of their habitat within the park. 

The southwestern corner of Brown County State Park was less affected by 
deer browsing than the main body of the park in both 1993 and 1994. The shared 
boundary with Yellowwood State Forest/Hoosier National Forest might have 
allowed increased hunting pressure in the southwestern corner of the park. 

No consistent significant difference for woody stem density (class 1) was 
found between the park and the State and National Forests. However, the high- 
er number of woody stems (class 2) outside the park when compared to both 
areas within the park suggests that the trees are successfully reproducing, but 
repeated browsing prevents seedlings from growing into the larger size classes. 

50 Ecology: Brown and Parker Vol. 106 (1997) 

Marquis (1974) found that browsing by white-tailed deer on the Allegheny Plateau 
in Pennsylvania prevented stems from growing beyond the seedling stage. His 
conclusions agree with those of Frelich and Lorimer (1985) and Anderson and 
Katz (1993), who both concluded that browsing by white-tailed deer was severe- 
ly affecting the reproduction and recruitment of eastern hemlock into larger size 

The large number of plant species still found within Brown County State 
Park in 1993 and 1994 indicates a potential for recovery of the plant communi- 
ties if deer populations are maintained at lower levels. Other researchers have 
also suggested that controlling deer numbers can effectively decrease browse 
damage (Anderson and Loucks, 1979; Frelich and Lorimer, 1985; Alverson, 
et ah, 1988; Anderson and Katz, 1993). Anderson and Loucks (1979) suggest 
that wildlife should be maintained at a level that would not be detrimental to 
important tree species. They also suggest that in areas where wildlife is having 
a detrimental effect, hunting regulations should be relaxed to prevent changes 
in tree species composition. Significant improvement in the understory could 
occur within a few years if deer numbers were maintained at lower levels (Ander- 
son and Loucks, 1979). However, a significant period of time will be needed 
before the forest can completely recover. Anderson and Katz (1993) suggest that 
the time needed for recovery after release from browsing pressure is directly pro- 
portional to the time the forest was subjected to this pressure. 

The limited removal of 392 deer (17 deer per square mile or 2.590 km 2 ) from 
the park in 1993 may have been responsible for the slight improvement in the 
condition of the understory vegetation. An increase in the number of plant species 
found per site occurred within the park in 1994. However, this increase in species 
number might be the result of deer reduction or annual variation. Before con- 
crete conclusions can be reached about the specific damage caused by deer, more 
data needs to be collected during subsequent growing seasons. Continued stud- 
ies of the understory vegetation could show not only differences in the degree 
of impact due to white-tailed deer browsing but also variation in the vegetation 
due to annual variability and the timing of sampling within the growing sea- 


The authors would like to thank the Indiana Department of Natural Resources 
for financial support. They would also like to thank Jim Eagleman and the staffs 
at Brown County State Park and Yellowwood State Forest for their assistance 
with this project. A special thanks to Amanda Mcintosh and Sally Weeks for aid- 
ing with data collection. 


Alverson, W.S., D.M. Waller, and S.L. Solheim. 1988. Forests too deer: Effects in northern Wisconsin. Cons. 
Biol. 2: 348-358. 

Anderson, R.C. and A.J. Katz. 1993. Recovery of browse-sensitive tree species following release from white- 
tailed deer (Odocoileus virginianus Zimmerman) browsing pressure. Biol. Cons. 63: 203-208. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 5 1 

and O.L. Loucks. 1979. White-tailed deer (Odocoileus virginianus) influence on structure and 

composition of Tsuga canadensis forests. J. Appl. Ecol. 16: 855-861. 

Balgooyen, C.P. and D.M. Waller. 1995. The use of Clintonia borealis and other indicators to gauge impacts 
of white-tailed deer on plant communities in northern Wisconsin, USA. Natur. Areas J. 15: 308-318. 

Brown County State Park Deer Study Committee. 1993. A report to the Natural Resources Commission. Indi- 
ana Dep. Natur. Res., Indianapolis, 12 pp. 

Brown, D.T. and G.J. Doucet. 1991. Temporal changes in winter diet selection by white-tailed deer in a north- 
ern deer yard. J. Wildlife Manage. 55: 361-376. 

Frelich, L.E. and C.G. Lorimer. 1985. Current and predicted long-term effects of deer browsing in hemlock 
forests in Michigan, USA. Biol. Cons. 34: 99-120. 

Gillingham, M.P. and F.L. Bunnell. 1989. Effects of learning on food selection and searching behavior of deer. 
Can. J. Zool. 67: 24-32. 

Homoya, M.A., D.B. Abrell, J.R. Aldrich, and T.W. Post. 1985. The natural regions of Indiana. Proc. Indiana 
Acad. Sci. 94: 245-268. 

Kohlman, S.G. and K.L. Risenhoover. 1994. Spatial and behavioral response of white-tailed deer to forage 
depletion. Can. J. Zool. 72: 506-513. 

Korschgen, L. 1962. Foods of Missouri deer, with some management implications. J. Wildlife Manage. 26: 

Marquis, D.A. 1974. The impact of deer browsing on Allegheny hardwood regeneration. U.S. Dep. Agr., U.S. 
Forest Serv. Paper NE-308, 8 pp. 

McArthur, C, C.T. Robbins, A.E. Hagerman, and T.A. Hanley. 1993. Diet selection by a ruminant generalist 
browser relative to plant chemistry. Can. J. Zool. 71: 2236-2243. 

McCaffery, K.R., J. Tranetzki, and J. Piechura, Jr. 1974. Summer foods of deer in northern Wisconsin. J. 
Wildlife Manage. 38(2): 215-219. 

Mumford, R. and J. Whitaker, Jr. 1982. Mammals of Indiana. Indiana Univ. Press, Bloomington, Indiana, 
537 pp. 

Nixon, C, M.W. McClain, and K.R. Russell. 1970. Deer food habits and range characteristics in Ohio. J. 
Wildlife Mgt. 34(4): 870-886. 

Parker, G.R. and J.E. Van Kley. 1993. Brown County State Park: Final report on vegetation damage by 
white-tailed deer. Unpubl. report. 

Smith, W.P 1991. Odocoileus virginianus. Mammalian Spec. 388: 1-13. 

Strole, T.A. and R.C. Anderson. 1992. White-tailed deer browsing: Species preferences and implications for 
central Illinois forests. Natur. Areas J. 12(3): 139-144. 

Van Kley, J.E. and G.R. Parker. 1993. An ecological classification system for the Central Hardwoods 
Region: The Hoosier National Forest, Indiana. Proc. 9th Central Hardwoods Conf., U.S. Dep. Agr., For- 
est Serv., North Central Forest Exp. Stat., Gen. Tech. Rep. NC-161, pp. 308-326. 

Westoby, M. 1974. An analysis of diet selection by large generalist herbivores. Amer. Natur. 108: 290-304. 

Proceedings of the Indiana Academy of Science 5 3 

(1997) Volume 106 p. 53-66 







Thomas P. Simon, Robert N. Jankowski, and Charles Morris 

Department of Biology 

1401 South U.S. 421 

Purdue University-North Central 

Westville, Indiana 46391-9528 

ABSTRACT: Four lakes located in the Indiana Dunes National Lakeshore 
in northwestern Indiana were evaluated for their physical and chemical lim- 
nology. The morphometric characteristics for each lake were measured as were 
chemical variables, including pH, conductivity and major ions, and nutrients. 
The study lakes were shallow depressions (z m = 1.17-3 m; z = 0.8-2.51) with 
steep slopes. Their shallow depths enabled dissolved oxygen to be distributed 
throughout the entire water column; thus, these lacustrine wetlands never strat- 
ify. The pH in these lakes was neutral to slightly alkaline (range = 7.20-10.02), 
while their conductivity ranged from 40 to 1339 juS/cm. The amount of total 
dissolved solids ranged from 48.6 to 67.0 mg/L. The oxidation-reduction poten- 
tial showed a stepwise progression in values ranging from -60 to 475 mv, and 
more than 90% of the measurements were in an oxidized state. The presence 
of an oxidized microzone above the sediment interface with the water column 
prevents metals and nutrients from autochthonous recycling. Total nitrogen 
levels were similar to those in mesoeutrophic lakes in northeastern and north- 
central Indiana, while total phosphorus (x = 0.121 mg/L) was an order of mag- 
nitude higher than in most of the lakes in north-central (x = 0.025 mg/L) and 
northeastern (x = 0.052 mg/L) Indiana. 

KEYWORDS: Bathymetry, conductivity and major ions, nutrients, pH, troph- 
ic status. 


Most regional studies of the chemical and physical limnology of glacial lakes 
have been carried out in Minnesota, Wisconsin, and Michigan (Heiskary. et 
a/., 1987; Omernik and Gallant, 1988). Glacial lake limnology along the dunes 
and nearshore of Lake Michigan has been devoted mainly to the recognition of 
environmental indicators and the identification of the aquatic biota (Simon, et 
a/., 1989; Simon and Moy, 1997; Simon and Stewart, in press). Little attention 
has been given to the physical and chemical attributes of riverine and depres- 
sional wetlands in northwestern Indiana. 

The greatest concentration of depressional wetlands in Indiana exists with- 
in the Indiana Dunes National Lakeshore (Beaty, et al, 1994). Knowledge of the 

54 Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

status and condition of these palustrine and lacustrine wetlands is limited. Many 
have never been surveyed. The collection of baseline data is necessary for deter- 
mining patterns in lake trophic status and for analyzing the physical attributes 
of other natural systems in northwestern Indiana. The morphologic features of 
these lakes were determined by climatic and edaphic factors that affect the chem- 
ical dynamics of the lake, which, in turn, shapes the biota within these ecosys- 

Early investigations of the physical and chemical environment in the nat- 
ural lakes of Indiana concentrated on the glacial lakes of northeastern Indiana. 
Classic studies of lake morphometries in Indiana include the State Geological 
Survey report on northern lakes (Blatchley and Ashley, 1901), the physical and 
biological studies of Lake Maxinkuckee (Evermann and Clark, 1920), and numer- 
ous lake studies conducted by the Indiana Lakes and Stream Survey (Scott, et 
al, 1928, 1938; Scott, 1931; Wohlschlag, 1950; Gerking, 1950; Ricker, 1955; 
Eberly, 1959; Mueller, 1964). The National Eutrophication Survey conducted 
by the U.S. Environmental Protection Agency in cooperation with the Indiana 
Department of Environmental Management (formerly a part of the Indiana State 
Board of Health) during the mid-1970s evaluated 27 lakes in Indiana (numer- 
ous individual reports published by U.S. Environmental Protection Agency, 
1976). These studies provided data on the chemical and physical characteris- 
tics of northern Indiana lakes and discussed their trophic status. The western- 
most lake surveyed during that study was Bass Lake in Starke County. 

Our objective was to describe the chemical and physical characteristics of 
four natural palustrine and lacustrine wetlands in northwestern Indiana. While 
many aspects of their biota can be determined without a knowledge of the phys- 
ical and chemical characteristics of these lakes, many of the indices of produc- 
tivity cannot be used without these data. This survey was designed to collect data 
on the physical and chemical characteristics of four typical palustrine and lacus- 
trine depressional wetlands in the Indiana Dunes National Lakeshore. In addi- 
tion to discussing the chemical and limnological characteristics of these lakes, 
our findings will be compared with data from other natural lakes in northern 
Indiana that were collected during the National Lake Eutrophication Survey. 


Description of the Study Area. The nearshore of Lake Michigan includes 
a variety of depressional wetlands, such as pannes, ponds, and lakes (Figure 1). 
Several distinct dune beach complexes were formed during the Pleistocene and 
Holocene Epochs when Lake Michigan was at higher levels than today (Lev- 
erett and Taylor, 1915; Bretz, 1951; Hansel, et aL, 1985). The area is part of a 
province referred to by various scientists as the Calumet Lacustrine Plain (Schnei- 
der, 1966), the Central Corn Belt Plain Ecoregion (Omernik and Gallant, 1988), 
or the Lake Michigan Border Section of the Northwestern Morainal Natural 
Region (Homoya, et al., 1985). This region is a mosaic of natural and human- 
impacted areas, including the Indiana Dunes National Lakeshore, the Indiana 

Vol. 106 (1997) Indiana Academy of Science 



Indiana Dunes 



Grand Calumet 

Figure 1 . A map of northwestern Indiana showing the location of the four lacustrine wet- 
lands studied. 

Dunes State Park, and the Clark and Pine Natural Area. To the west lies one of 
the most industrialized steel and petrochemical areas in the United States (Moore 
and Trusty, 1977; Simon, et ai, 1989). The four depressional wetlands discussed 
in this paper are found in the Indiana Dunes National Lakeshore. 

Regional Climate and Hydrology. The prevailing climate in northwest- 
ern Indiana is temperate continental modified by Lake Michigan so that the cli- 
mate can take on semi-marine characteristics. The mean annual temperature is 
10° C. Average annual precipitation at Gary (the largest nearby city) is about 
907.5 mm; normal seasonal precipitation averages 145 mm in the winter, 252.5 
mm in the spring, 285 mm in the summer, and 225 mm in the fall (National 
Oceanic and Atmospheric Administration, 1982). Total monthly rainfall is more 
variable during warm months that during cold months. The total annual precip- 
itation between 195 1-1980 ranged from about 575 mm to nearly 1250 mm. Annu- 
al snowfall varies due to the lake effect. Annual average snowfall is 875 mm at 
Gary with the predominant snow season from November to March. Due to the 
proximity of the study wetlands to Lake Michigan, early frosts and unusually 
late spring frosts may be delayed by 2-3 weeks. The coldest month (January) has 
an average normal monthly temperature of -5.1° C; the average normal month- 
ly temperature during the warmest month (July) is 22.9° C (Beaty, et al.. 


Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

Middle Grand Calumet Lagoon 

East Grand Calumet Lagoon 



Figure 2. Bathymetric contour maps of the four study lakes: a = Middle Grand 
Calumet Lagoon; b = East Grand Calumet Lagoon; c = Long Lake; and d = Little 
Lake. The depth contour line intervals are 0.5 m. 

The ponds of West Beach and Miller Woods comprise the last remnant, 
depressional wetlands in northwestern Indiana along the Lake Michigan dunes 
in the Indiana Dunes National Lakeshore. These ponds are remnants of geolog- 
ical Lake Chicago that were created by the lowering of lake levels and the shift- 
ing of sand dunes (Moore and Trusty, 1977). Long Lake is a large, dunal pond 
located southeast of Miller Woods in the West Beach Segment. The Grand Calumet 
Lagoons are riverine wetlands derived from the Grand Calumet River. The 
Lagoons are the former mouth of the river. They are separated from the river 
by a covered culvert on adjacent industrial property. The Lagoons are divided 
into three segments of relatively equal area — the East, Middle, and West Lagoons. 
This study was confined to the Middle and East Lagoons, the "least disturbed" 
ponds in the Indiana Dunes National Lakeshore and the City of Gary's Marquette 
Park. Little Lake is a former portion of Cowles Bog that was separated from 
the bog by a levee built for an electric utility substation. 

Bathymetry, Morphometry, and Chemical Limnology. The bathymetric 
contour maps of the four lakes (Figure 2) were prepared by tracing the shore- 
lines from perimeter maps obtained from the analysis of aerial photographs 
and topographic maps (7.5 minute U.S. Geological Survey maps). The size and 
complexity of each lake determined the number of transects needed to obtain an 

Vol. 106 (1997) Indiana Academy of Science 57 

appropriate bathymetric profile. Little Lake, which is the smallest of the lakes 
at about 6 ha, was mapped at 20 m transect intervals; Long Lake, which is about 
25.9 ha, was mapped using a 30 m interval; and the Middle (30.3 ha) and East 
Grand Calumet Lagoons (5.1 ha), the largest of the lakes, were studied using 
50 m transect intervals. Intervals were measured by determining the perpendic- 
ular intersection of adjacent shorelines and then measuring the distance between 
points using a calibrated line (Cole, 1994). Points were then marked where 
they would be visible from the water using flagging tape. Depths were record- 
ed to the nearest 0. 1 m at discrete points along each transect from a boat. A Hum- 
mingbird 3-D depth-finder was used to sample nonwadeable areas, while depths 
were recorded in centimeters using a Philadelphia rod in wadeable areas. Coor- 
dinates were then plotted using Grass 4.0, and a detailed bathymetric map was 
drawn using Winsurf 5.0 (Golden Software, 1994). 

Morphometric parameters were calculated from these maps, following the 
procedures of Lind (1985) and Wetzel and Likens (1979). Surface morphomet- 
ric measures included maximum length (1), maximum width (b), mean width (b), 
surface area (A), shoreline length (L), shoreline breath, and the shoreline devel- 
opment index (D L ). Subsurface morphometries included volume (V), maximum 
depth (z m ), mean depth (z), relative depth (z r ), and basin slope. Morphoedaphic 
index calculations followed Ryder, et at. (1974). Temperature and dissolved oxy- 
gen were measured monthly between February 1992 to August 1997 at near bot- 
tom stations at the approximate center of each lake. 

The four lakes were too shallow to develop a stratified profile. Most parts 
of each lake are less than 2 m in maximum depth. A digital meter (Dow 
Corning Inc., Pocket Meter M90) was used to measure dissolved oxygen 
(DO; 0.0 to 20.00 ± 0.1 mg/L), temperature (-0.5° to 100° ± 0.1° C), pH 
(0 to 14 ± 0.1 SU), specific conductance (0.0 to 1999 ± 1 ^S), and total dissolved 
solids (TDS; 0.0 to 1000 ± 0.1 mg/L). The oxidation-reduction potential (E h ) 
was measured using a digital meter (LaMotte, Inc., ORPTestr, -200 to 1 100 ± 5 
mv). Dissolved oxygen was calibrated using Winkler titration (American Pub- 
lic Health Association, 1989). Water samples were taken at the deepest level of 
the water column (near bottom) along predetermined transects using a Kem- 
merer water bottle sampler. Water samples from each lake were analyzed at the 
Indiana Dunes National Lakeshore Aquatic Laboratory for nitrite, nitrate, ammo- 
nia, hardness, alkalinity, and reactive phosphorous using the appropriate meth- 
ods (American Public Health Association, 1989). 


Origin. The analysis of aerial photographs of the Grand Calumet Lagoons 
suggests that these water bodies originated following closure of the river's mouth 
by dune movement. The East and Middle Lagoons were extensive backwaters 
at the turn of the century when dune movement separated them from the Grand 
Calumet River. These two ponds are separated by a large lateral dune that aris- 
es in the foredune ridge near Lake Michigan, suggesting that the two lakes along 

58 Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

Table 1 . Morphometric characteristics of four lakes from northwestern Indiana. 

East Grand 

Middle Grand 



Calumet Lagoon 

Calumet Lagoon 



Maximum length (1) 

1730 m 

1290 m 

437 m 

138 m 

Maximum depth (z m ) 

3.0 m 

3.0 m 

1.2 m 

1.8 m 

Maximum width (b) 

330 m 

370 m 

273 m 

378 m 

Perimeter (L) 

4250 m 

4060 m 


4622 m 

Shoreline development (D L ) 

2.3 m 

2.7 m 

1.7 m 

2.5 m 

Surface area (A) 

264,090 m 2 

318,181.8 m 2 

63,029 m 2 

272,000 m 2 

Volume (V) 

663,634 m 3 

633,181.8 m 3 

5,044 m 3 

304,000 m 3 

Mean width (b) 

152.7 m 

246.7 m 

144 m 

197 m 

Mean depth (z) 

2.5 m 

2.0 m 

0.8 m 


Relative depth (z r ) 

0.5 m 

1.5 m 

0.6 m 

0.3 m 

Volume development (D v ) 

> 1 

> 1 

> 1 

> 1 

Basin slope (M) 





Morphoedaphic index (MEI) 





with the Grand Calumet River were once part of a single, continuous watershed. 
Long Lake is an 8,000-year-old successional lake formed by the regression of 
Lake Michigan (R. Whitman, pers. comm.). The eastern end of Long Lake has 
been filled in, creating a separation between Ogden Dunes and the extensive wet- 
lands previously occurring along the Little Calumet River. Long Lake is exten- 
sively vegetated and numerous beds of pondweeds (e.g., Potemogeton) and water 
lilies {Nuphar and Nymphea) have created a vegetated mat that is filling the shal- 
low lake. An increase in nutrients has caused the degradation of this fragile 
system. Little Lake is the former southwestern corner of Cowles Bog, a large 
palustrine wetland filled with Typha. The levee along its northern shore sepa- 
rates Little Lake from Cowles Bog. To accommodate an electric substation on 
its northwestern shore, Little Lake was deepened and a narrow access corridor 
to an electric tower was built into the water. A natural area on the southeastern 
shore has fallen woody debris, emergent aquatic macrophytes, and submergent 
stands of Ceratophyllum demersum and Myriophyllum spicatum. The diver- 
sion of groundwater has flooded the area between the South Shore railroad tracks 
and the levee. 

Bathymetry and Morphometries. Bathy metric maps of the four lakes are 
shown in Figure 2, and their morphometries are listed in Table 1 . These lakes 
have surface to volume ratios that are very low, a feature characteristic of the 
natural lakes found in northwestern Indiana. The deepest lake is the East Grand 
Calumet Lagoon, whose mean depth is 2.51 m. The shallowest lake was Little 
Lake, whose mean depth was 1.17 m. The basin slope (M) of these lakes 
ranges from 0.0184 to 0.027, confirming their shallow depths when compared 

Vol. 106 (1997) Indiana Academy of Science 59 

to their surface area. The shoreline development index (D L ) also shows little vari- 
ation; the index ranges from 1.703 to 2.73 (Table 1). These values of the shore- 
line development index are consistent with those from most lakes that develop 
increased littoral regions (Wetzel, 1983). The lakes trend east to west in latitu- 
dinal profile, they are elongate, and they have irregular shorelines. 

All four depressional wetlands lack major inflows. All four lakes are sub- 
ject to atmospheric inputs primarily as a result of air emissions from industrial 
sources. The majority of the allochthanous input into Middle and East Grand 
Calumet Lagoon is probably the result of urban and residential development, 
which generates a higher volume of runoff because of an increase in the area 
covered by an impervious surface. Little Lake receives leaf litter from the sur- 
rounding forest. Inputs of particulate matter in Long Lake are the result of air- 
suspended material from the adjacent dunes. The shallow depth of all these lakes 
contributes to their eutrophic condition because dissolved oxygen and light pen- 
etrates to the benthic region despite the presence of suspended solids in the water. 

Temperature and Oxygen. The four study lakes do not thermally stratify 
during either summer or winter. Dissolved oxygen is present in the entire water 
column, and a permanent oxidized microzone is present (Table 2). The mean dis- 
solved oxygen level for the Middle Grand Calumet Lagoon was approximately 
62% saturation, while the other three lakes had mean dissolved oxygen levels 
between 82% to 88% saturation. The amount of dissolved oxygen is strongly 
linked to temperature. The lowest amount of dissolved oxygen is detected when 
the temperatures are warmest (0.81 to 0.83 mg/L in Long and Little Lakes, respec- 
tively). Supersaturated dissolved oxygen values can occur because of the large 
aquatic macrophyte population that grows in all portions of the lake. Extreme 
diel fluctuations may occur because nocturnal respiration by these large plant 
beds can cause an oxygen deficit. 

pH. The pH was highly variable for lakes located in an area of relatively 
homogeneous geologic and edaphic conditions. The pH ranged from 6.70 to 
10.02 (Table 2). All four lakes are found along the edge of Lake Michigan and 
accept drainage from Oakville Maumee-Brems Soils (Furr, 1981). The dune, 
beach, and lacustrine silts, sands, and gravel form a thin but laterally extensive 
surficial aquifer. The sediments of Little and Long Lake arose from drained Adri- 
an muck, while the sediments of East and Middle Grand Calumet Lagoon are 
Oakville fine sand soils with slopes of 18% to 40% (Furr, 1981). Several sites 
in East Grand Calumet Lagoon had pH values above 10 SU. Locally elevated 
pH values may be the result of isolated, narrow belts of slag rock that were buried 
after the dunes were mined (R.D. Kovach, U.S. Environmental Protection Agency, 
pers. comm.). Little Lake and the East Grand Calumet Lagoon consistently had 
the greatest variation in pH (Table 2). 

In general, fresh water within the study area is not sensitive to acidification 
because the water is well buffered (Fenelon and Watson, 1993; Willoughby, 1995: 
Duwelius, et al, 1996). Fenelon and Watson (1993) reported median pH val- 
ues from groundwater well samples of about 7.3; four extreme pH values were 


Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

Table 2. Mean and standard deviation for monthly field and water chemistry data col- 
lected from February 1992 to August 1997 at four lacustrine wetlands in northwestern 
Indiana. The range is given in parentheses. 

East Grand 

Middle Grand 




Calumet Lagoon 

Calumet Lagoon 



Alkalinity (mg/L as CaC0 3 ) 

128 ±37.5 


139 ±50.0 

96 ± 24.4 




Hardness (mg/L as CaC0 3 ) 

204 ± 39.5 


194 ±47.6 

132 ±35.5 




pH (SU) 

8.2 ± 0.46 

8.34 ±0.10 

8.1 ±0.67 

8.0 ±0.38 





Temperature (° C) 

5.3 ±0.22 

5.0 ±0.15 

8.1 ±5.8 

5.8 ±0.38 





Chloride (mg/L) 

110 ±35.6 


74 ±25.61 

30 ± 19.45 




Specific Conductance 

609 ±213.6 

131.5 ±0.6 

491 ± 195.8 

248 ±62.1 






Dissolved Oxygen (mg/L) 

9.02 ± 3.03 

4.50 ± 0.07 

8.52 ±3.08 

8.00 ± 3.42 





Dissolved Oxygen 

88 ±26.14 

62 ±10.45 

87 ± 27.36 

82 ± 27.86 

(% saturation) 





NH, (mg/L) 

0.13 ±0.15 

0.14 ±0.28 

0.19 ±0.35 

0.15 ±1.87 





Nitrate (mg/L) 

0.40 ± 0.60 


0.3 ± 0.62 

0.3 ±0.61 




Nitrite (mg/L) 

0.003 ±0.124 


0.001 ±0.01 

0.001 ±0.002 




Reactive Phosphorus 

0.030 ± 0.054 


0.05 ± 0.05 

0.02 ± 0.03 





Total Phosphorus (mg/L) 

0.135 ±0.09 


0.12 ±0.26 

0.075 ± 0.08 





163 ± 115.1 

152.5 ± 143.4 

375 ±141.4 

235 ± 82.5 

Potential (E h ; mv) 





Total Dissolved Solids 

66.8 ± 7.49 

65.85 ±0.61 

63.5 ± 0.07 

59.5 ± 0.96 






Vol. 106 (1997) Indiana Academy of Science 61 

reported from wells screened in slag. Slag is dominantly alkaline-earth- silicate 
glass, and water in contact with it has an elevated pH. Duwelius, et al. (1996) 
conducted a more extensive survey of the groundwater in area wells and found 
that the pH ranged from 5.3 to 1 2. 1 (median = 7.2). Half of the 1 1 8 samples from 
wells had pH values between 6.9 and 7.6. For most of the groundwater in the 
United States, pH ranges from about 6 to 8.5 (Hem, 1985). Duwelius, etal. (1996) 
found that samples with an elevated alkaline pH were usually from shallow 
groundwater wells in contact with slag or industrial waste. 

Conductivity and Major Ions. Specific conductance is a measure of the 
ability of a substance to conduct electricity across a unit length at a specific tem- 
perature. Dissolved substances increase the conductivity of water; measurements 
of specific conductance provide an indication of the amount of dissolved sub- 
stances in water (Hem, 1985). The specific conductance of pure water is low, 
less than 10 jU,S/cm (Hem, 1985). In general, the surface waters of our study had 
moderate conductance. Conductivity was consistent among wetlands and ranged 
from 40 to 1,339 /xS/cm (mean = 369.9 /xS/cm; Table 2), which was consider- 
ably lower than the conductivity reported for groundwater wells (106-5,980 
jLtS/cm; median = 828 /u,S/cm). Half of the 125 well samples had specific con- 
ductance values between 450 and 1,540 /LtS/cm (Duwelius, etal, 1996). The East 
Grand Calumet Lagoon had the highest conductivity of the four lakes studied. 

Alkalinity measures the capacity of a solution to neutralize acids (Hem, 
1985). In this study, alkalinities ranged from 44 to 330 mg/L as calcium car- 
bonate (mean =121 mg/L). Duwelius, et al (1996) recorded alkalinities from 
groundwater wells in northwestern Indiana that ranged from 24.9 to 1,260 mg/L 
as calcium carbonate (median = 249 mg/L). Little Lake had the lowest alkalin- 
ity (Table 2). Acidity measures a solution's capacity to neutralize bases. Acidi- 
ty was not detected in any sample. 

Water is considered very hard when values exceed 1 80 mg/L as calcium car- 
bonate. All of the lakes have very hard water (mean = 198.7 mg/L) with the 
exception of Long Lake (mean = 132 mg/L; Table 2). Fenelon and Watson (1993) 
found that non-contaminated "natural" groundwater samples from the Indiana 
Dunes National Lakeshore never exceeded 200 mg/L. Groundwater samples col- 
lected from areas adjacent to the heavily industrialized areas surrounding the 
Grand Calumet River ranged from 400 to 500 mg/L, 

Freeze and Cherry (1979) placed groundwater samples into two categories 
based on total dissolved solids (fresh and brackish water). Fresh water general- 
ly contains less than 1,000 mg/L of dissolved solids. Water having between 1.000 
and 10,000 mg/L of dissolved solids is called brackish (Freeze and Cherry. 1979). 
In general, the surface waters in this study represented fresh water. The amount 
of dissolved solids was similar in each lake and ranged from 48.6 to 67.0 mg/L 
(mean = 63.9 mg/L; Table 2). Duwelius, et al (1996) found that groundwater 
wells had dissolved solid concentrations that ranged from 95 mg/L to 6,780 mg/L; 
the median concentration was 674 mg/L. A comparison of dissolved solid con- 
centrations between paired shallow and deep wells showed that the highest 
values come from shallow wells. 

62 Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

The oxidation-reduction potential (E h ) of water is an index of the exchange 
activity of electrons among elements in solution. E h measures the electric poten- 
tial, using the potential of a hydrogen electrode as a reference point of zero. A 
positive potential indicates oxidizing conditions in the water; a negative poten- 
tial indicates reducing conditions, which determines the valence states of met- 
als (Hem, 1985). The oxidation-reduction potential at the study sites ranged in 
a stepwise progression from -60.0 to 475 mv (mean = 231.4 mv). Duwelius, et 
al. (1996) found that groundwater from wells was generally reducing (75%), and 
E h ranged from -446 to 159 mv (median = -64.5 mv). Reducing conditions 
increased with increasing well depth. With the exception of the East and Mid- 
dle Grand Calumet Lagoon, few of our study sites ever showed reducing con- 
ditions; more than 90% of the sites were in an oxidized condition. 

Chloride is the dominant anion in water, and its concentration ranged from 
2.0 to 238 mg/L (mean = 71.3 mg/L; Table 2) in the lakes studied. Duwelius, et 
al. (1996) found that the concentration of chloride in groundwater wells ranged 
from 1 .4 mg/L to 2,600 mg/L (median = 37.8 mg/L). The wells that had the high- 
est concentrations of chloride (greater than 1,000 mg/L) were shallow (less than 
4.5 m deep) and were found in areas containing fill near interstate highways. 
High chloride concentrations indicate contamination by the fill materials and 
deicing salts. The Secondary Maximum Contaminant Levels are suggested con- 
centration limits for substances in drinking water that do not result in adverse 
health effects but may limit the use of the water because of unpleasant taste, odor, 
or color. The suggested limit for chloride is 250 mg/L, a value that was not exceed- 
ed in the study lakes. 

Nutrients. The concentrations of nitrate plus nitrite, ammonia, and reactive 
and total phosphorus were determined. Nitrogen concentrations were generally 
low; N0 3 , N0 2 , and NH 3 occurred in concentrations of less than 1 .0 mg/L. Total 
phosphorus levels were high (mean = 0.100 mg/L). The surface waters of the 
study lakes all had high concentrations of nutrients. 

Ammonia levels ranged from 0.01 to 0.78 mg/L (mean = 0.15 mg/L; Table 
2). This range was lower than that reported from groundwater well samples (0.1- 
96 mg/L; median = 0.50 mg/L) by Duwelius, et al. (1996). Fenelon and Wat- 
son (1993) found ammonia concentrations at the Indiana Dunes National Lakeshore 
to range from 0.05 to 0.20 mg/L. Long and Little Lakes had the highest mean 
ammonia concentrations. Their mean values were over twice as high as the 
values observed in the Grand Calumet Lagoons. Duwelius, et al. (1996) report- 
ed that half of the 125 samples from groundwater wells had ammonia (nitrogen) 
concentrations less than the detection level (< 0.01 mg/L). 

The concentration of nitrate plus nitrite was low, ranging from 0.001 to 2.5 
mg/L (mean = 0.38 mg/L; Table 2). These concentrations were higher than those 
found by Fenelon and Watson (1993). Their values for the concentration of nitrate 
plus nitrite in groundwater samples from the Indiana Dunes National Lakeshore 
ranged from below detection level (< 0.01 mg/L) to 0.02 mg/L and were with- 
in the range reported for natural groundwater samples (range = 0.02 to 0.96 mg/L; 
median = 0.06) in northwestern Indiana. 

Vol. 106 (1997) Indiana Academy of Science 


Table 3. A comparison of select physical, spring chemical, and trophic status variables 
for a variety of natural lakes in northern Indiana (N = nitrogen and P = phosphorus). 

Physical Variables (m) 

Chemical Variables 




N-Total P-Total 



(m 3 ) 


Depth (z) 

(mg/L) (mg/L) 



Middle Grand Calumet Lagoon 

0.6 x 10 6 






East Grand Calumet Lagoon 

0.6 x 10 6 






Little Lake (Porter) 

0.05 x 10 6 






Long Lake (Porter) 

0.3 x 10 6 







James Lake (Kosciusko) 

9.348 x 10 6 






Tippecanoe Lake (Kosciusko) 

35.143 xlO 6 






Lake Wawasee (Kosciusko) 

82.946 x 10" 






Webster Lake (Kosciusko) 

4.977 x 10 6 






Winona Lake (Kosciusko) 

20.657 x 10 6 






Lake Maxinkuckee (Marshall) 

55.042 x 10 6 







Dallas Lake (LaGrange) 

12.303 x 10 6 






Olin Lake (LaGrange) 

4.914 x 10 6 






Oliver Lake (LaGrange) 

18.300 xlO 6 






Sylvan Lake (Noble) 

10.879 x 10 6 






Crooked Lake (Steuben) 

19.825 x 10 6 






Hamilton Lake (Steuben) 

20.475 x 10 6 






Long Lake (Steuben) 

1.887 xlO 5 






Marsh Lake (Steuben) 

1.403 x 10 6 






Lake James (Steuben) 

30.514 x 10 6 






Pigeon Lake (Steuben) 

1.143 x 10 6 






The total and reactive phosphorus levels in the lakes of northwestern Indi- 
ana were higher than those in lakes from the remainder of northern Indiana (Table 
3). The total phosphorus concentrations in the lakes of northwestern Indiana were 
an order of magnitude higher than those for other lakes measured during the 
National Eutrophication Survey (U.S. Environmental Protection Agency, 1976). 
Total phosphorus concentrations in the four study lakes ranged from 0.01 to 1.58 
mg/L (mean = 0.1 mg/L; Table 2). The reactive phosphorus in the four study 
lakes was about 25% of total phosphorous with the exception of East Grand 
Calumet Lagoon, which was less than 25% (Table 2). 

Comparison to Natural Lakes. Ponds and lakes along the nearshore of 
Lake Michigan are shallow, eutrophic depressions that have steep littoral slopes 
formed by the erosion and movement of sand dunes. The lakes of northwestern 
Indiana are significantly different from other natural lakes in northern Indiana. 
The lakes of northwestern Indiana have an average maximum depth of 2.3 m in 
contrast to average lake depths of 9.38 m and 7.45 m for north-central and north- 
eastern Indiana, respectively (Table 3). The deeper lakes of central and eastern 
Indiana stratify and develop thermoclines. The lakes of northwestern Indiana 

64 Ecology: Simon, Jankowski, and Morris Vol. 106 (1997) 

never stratify because they are not deep enough. These lakes rarely attain aver- 
age depths greater than 2.0 m (mean = 1.6 m; Table 3). The maximum depth of 
north-central Indiana lakes ranged from 13.7 (Webster Lake, Kosciusko Coun- 
ty) to 37.5 m (Tippecanoe Lake, Kosciusko County). Northeastern Indiana lakes 
ranged from 9.7 m (Long Lake, Steuben County) to 29.3 m (Dallas Lake, LaGrange 
County) in maximum depth. 

Most of the large lakes of northern Indiana are either eutrophic or mesoeu- 
trophic, based on the trophic index developed by the Indiana Department of Envi- 
ronmental Management (1986). The index uses fifteen measurements (based on 
data from 307 samples collected during the mid-1970s) to characterize the troph- 
ic status of inland Indiana lakes. The measurements include broad categories 
of nutrients, dissolved oxygen, light penetration, total plankton, dominance of 
blue-green algae, and the abundance of cells in vertical tows from the thermo- 
cline and from the littoral zone. 

The lakes studied in the Indiana Dunes National Lakeshore are eutrophic or 
hypereutrophic, exhibit rapid cycling of nutrients, and show predictable crash- 
es in dissolved oxygen during diel cycling. These lakes also had low levels of 
total nitrogen but high levels of total phosphorus (Table 3). Average total nitro- 
gen for lakes in north-central Indiana ranged from 0.757 mg/L (Lake Maxinkuc- 
kee, Marshall County) to 2.159 mg/L (Winona Lake, Kosciusko County), 
while the lakes in northeastern Indiana had higher concentrations, ranging 
from 0.771 mg/L (Lake James, Steuben County) to 4.087 mg/L (Pigeon Lake, 
Steuben County). Total phosphorous for lakes in north-central Indiana ranged 
from 0.007 mg/L (Lake Wawasee, Kosciusko County) to 0.041 mg/L (Winona 
Lake, Kosciusko County), while total phosphorous in northeastern lakes ranged 
from 0.011 mg/L (Olin Lake, LaGrange County) to 0.166 mg/L (Sylvan Lake, 
Noble County). The total phosphorous in northwestern Indiana lakes is similar 
to the amount in mesoeutrophic lakes in north-central and northeastern Indiana 
(Table 3). The presence of an oxidized microzone in the lakes of northwestern 
Indiana limits autochthonous cycling of nutrients causing permanent loss of phos- 
phorus and nitrogen to the sediments. 


The authors wish to acknowledge the assistance of Anita Arends and Bob 
Daum, National Park Service, Indiana Dunes National Lakeshore, and P. Michael 
Stewart, Richard Whitman, and Jason Butcher, U.S. Geological Survey, Lake 
Michigan Ecology Station, for professional courtesies including use of labora- 
tory space as well as access to maps and other unpublished documents. Mary 
Bennet assisted in the use of Grass 4.0 for file management and Winsurf 4.0 
for plotting Figure 2. Lou Brenan, Indiana Dunes National Lakeshore, provid- 
ed an electronic version of Figure 1 that was modified by the senior author, and 
Charlie Morris prepared Figure 2. The authors appreciate the assistance of Eric 
Garza and Lora Hebert in the field. This is Publication Number 2 of the Indi- 
ana Biological Survey, Aquatic Research Center, Northwest Regional Center. 

Vol. 106 (1997) Indiana Academy of Science 65 


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Central Corn Belt Plain. U.S. Environ. Prot. Agency, Region 5, Environ. Sci. Div., Monitoring Quality 

Assurance Branch, Ambient Monitoring Sect., Chicago, EPA 905-91-025, 1 18 pp. 
, G. Bright, J. Rud, and J.R. Stahl. 1989. Water quality characterization of the Grand Calumet 

River basin using the index of biotic integrity. Proc. Indiana Acad. Sci. 98: 257-265. 
and PB. Moy. 1997. Historical, present, and future trends and condition offish communities in 

the Grand Calumet River and Indiana Harbor Canal. U.S. Army Corps Eng., Chicago District, Chica- 
go, Illinois, 27 pp. 
and P.M. Stewart, in press. Structure and function offish communities in the lower Lake Michi- 

gan drainage with emphasis on restoration of native fish communities. Natur. Areas J. 

Stahl, J.B. 1959. The developmental history of the chironomid and Chaoborus faunas of Myers Lake. Invest. 
Indiana Lakes Streams 5: 47-102. 

U.S. Environmental Protection Agency. 1976. Report on individual lakes from Indiana. National Eutrophica- 
tion Survey. Working Paper Series Numbers 325, 326, 328, 330, 33 1 , 332, 333,335,338, 339, 340, 34 1 , 
342, 344, 345, 348. U.S. Environ. Prot. Agency, Corvallis Environ. Res. Lab., Corvallis, Oregon. 

Watson, L.R., R.J. Shedlock, K.J. Banaszak, L.D. Arihood, and PK. Doss. 1989. Preliminary analysis of the 
shallow ground-water system in the vicinity of the Grand Calumet River-Indiana Harbor Canal, north- 
western Indiana. U.S. Geol. Surv. Open File Rep. 88-49, 45 pp. 

Wetzel, R. and G.E. Likens. 1979. Limnological analyses. W.B. Saunders, Philadelphia, Pennsylvania, 357 

Willoughby, T.C. 1995. Quality of wet deposition in the Grand Calumet River watershed, northwestern Indi- 
ana, June 30, 1992-August 31, 1993. U.S. Geol. Surv. Water Res. Invest. Rep. 95-4172, 55 pp. 

Wohlschlag, D.E. 1950. Vegetation and invertebrate life in a marl lake. Invest. Indiana Lakes Streams 3: 

Proceedings of the Indiana Academy of Science 67 

(1997) Volume 106 p. 67-77 






Ronald A. Weiss 

Chipper Woods Bird Observatory 

10329 North New Jersey Street 

Indianapolis, Indiana 46280 

ABSTRACT: The house finch (Carpodacus mexicanus), first released in New 
York in 1940, has expanded its range throughout North America, making its 
first appearance in Indiana in the mid-1970s. The finch's first appearance in 
an annual Indiana Audubon Society bird count occurred in 1980, when two 
individuals were counted in Porter County during the Indiana Audubon Soci- 
ety Big May Day Bird Count. Five individuals were counted in the 1981 sum- 
mer count, and a single individual was counted in the 1981 Indiana Audubon 
Society Christmas count. Since then, the annual Indiana Audubon Society bird 
counts have shown a dramatic increase in house finch numbers in Indiana. 
Trend analysis of the Indiana Audubon Society May, summer, and Christ- 
mas counts reveal that the Christmas counts are increasing at the greatest rate, 
followed by the May counts, and then the summer counts. Moving average 
plots of house finch counts show that count trends have been exponential with 
interruptions. House finch populations may be approaching their upper limit 
in Indiana. 

KEYWORDS: Bird populations, Carpodacus mexicanus, Christmas bird 
counts, house finch, Indiana. 


The house finch {Carpodacus mexicanus), a Western species first released 
in New York in 1940 (Hill, 1993), has expanded westward, making sporadic 
appearances in Indiana from the mid-1970s to the early 1980s (Clay and Clay, 
1981; Heller and Wise, 1982; Hill, 1993; Gill, 1984; Wiggins, 1987). The house 
finch was first recorded in the 1980 Indiana Audubon Society May count, 
when two individuals were observed in Porter County in northwestern Indiana 
(Hopkins, 1980). Five individuals were reported in the 1981 summer count (Jack- 
son, 1983), and a single house finch was reported in the 1981 Christmas bird 
count (Mason and Mason, 1982). The number of house finches reported in sub- 
sequent counts has increased dramatically (Figure 1). The first breeding record 
for the house finch in Indiana occurred in Adams County in 1981 (Heller and 
Wise, 1982). 

Studies of the house finch in Indiana have focused on the Christmas count 
data and the impact of the house finch on house sparrow (Wise and Walls. 1988: 
Hamilton and Wise, 1991), American goldfinch, and purple finch populations 


Ecology: Weiss 

Vol. 106 (1997) 

8000 i 




| 3000 




Christmas Count 
May Count 
Summer Count 

© T- (VJ 


ff> ff» C"> 

r- CM 

Figure 1. The Indiana Audubon Society house finch counts from 1980 to 1995. 

(Hamilton and Wise, 1991). House finches appear to negatively impact house 
sparrow populations in the northeastern United States (Kricher, 1983). In Indi- 
ana, however, house sparrow populations may have declined prior to the 
arrival of the house finch (Hamilton and Wise, 1991). 

What trends are reflected in the Indiana Audubon Society May and summer 
house finch counts? How do the Indiana Audubon Society May, summer, and 
Christmas count trends for this species compare with each other? Can these trends 
be used to predict the size of the future house finch population? 


House finch count data collected annually by volunteers from the Indiana 
Audubon Society in the Christmas, May, and summer counts were compiled 
from records published in the Indiana Audubon Quarterly from 1980 to 1996. 
The data collection protocols for each of these counts differed, making their com- 
parison difficult. The May and summer counts were conducted county-by-coun- 
ty. The Christmas count, on the other hand, is not county based but is conducted 
within a 15 mile (24 km) diameter circle around an established center. Unlike 
the May and summer counts, at least 8 hours must be spent at each Christmas 
count site. Christmas counts are taken over a two week period. The May count 
occurs on the second Saturday in May. 

Vol. 106 (1997) Indiana Academy of Science 






50 ■■ 


Christmas Count Index 
May Count Index 
Summer Count Index 















r- co 

00 00 

en en 









<3" LD 

cn en 




The Indiana Audubon 

Society house finch count indices from 1980 to 1995. 

To compare data for the Christmas, May, and summer counts, a species 
detectability index was calculated for each respective count by dividing the total 
number of house finches observed in that count by the number of counties or 
Christmas counts participating each year. An index based on the number of field 
hours or the number of party hours could have been used, but those data were 
not complete for the period under study. The number of counties reporting 
each year reflects the total field effort and, by its nature, minimizes the variability 
in individual observer skill and the effort inherent in each respective count. 
The Christmas count trend, generated using the number of counts, is similar to 
the curve obtained by Hamilton and Wise ( 199 1 ) using the number of Christmas 
count party hours. 

The raw data for the Christmas, May, and summer counts were used to cal- 
culate a linear regression equation for each respective count (Figure 2). Linear 
regression analysis was also carried out using the index for each of the three 
counts (Figure 4). A moving average was calculated for each count based on the 
following formula and then plotted (Figures 5-7): 


_ 1 

Jv lAr " /+1 


Ecology: Weiss 

Vol. 106(1997) 










Figure 3. Indiana house finch count trends from 1980 to 1995. 

where N, the number of periods included in the moving average, is 3, A t _ i+l is the 
actual value at time J, and F (t+l) is the forecasted value at time J. Data analysis 
was carried out using Microsoft Excel on a Macintosh Performa 6300CD. 


House finch counts in Indiana have increased dramatically since the house 
finch was first reported in the Indiana Audubon Society's Christmas, May, and 
summer bird counts (Table 1). Plots of the count data from 1980 to 1995 demon- 
strate that the Christmas counts increased the most with the May counts and sum- 
mer counts following in that order (Figure 1). The regression equation (Figure 
3) calculated using the Christmas data had a slope of 644 (y = 644x - 1728), more 
than twice that for the May data, whose slope was 250 (y = 250x - 921), and 
more than four times that of the summer data, whose slope was 156 (y = 156x - 

Plots of the count indices for the period 1980-1995 (Figure 2) were similar 
to those for the raw data (Figure 1). Regression equations for the count indices 
for Christmas, May, and summer were calculated and plotted (Figure 4). The 
Christmas count index had a slope of 17 (y = 17x - 45), more than twice that of 
the May count index, whose slope was 6 (y = 6x - 16), and nearly five times that 
of the summer count index, whose slope of 3.6 (y = 3.6x - 11). Both the raw 

Vol. 106 (1997) Indiana Academy of Science 



£ 200 



Christmas Index 
Christmas Trend 
May Index 
May Trend 
Summer Index 
Summer Trend 


Figure 4. Indiana house finch index trends from 1980 to 1996. 

Table 1 . Indiana Audubon Society house finch count data. 

1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 

Christmas counts 

Number of counts 
House finch count 
Christmas Index 1 

May counts 

35 34 40 39 40 42 42 40 39 31 30 35 37 42 39 
1 56 179 178 766 1259 2942 4357 3600 3226 4777 7769 6842 7474 
0.03 1.4 4.6 4.5 18 30 74 112 116 108 136 210 163 192 

Number of Counties 

reporting 40 39 41 42 

House finch count 2 7 13 65 

Index 2 0.05 0.18 0.32 1.5 

Summer counts 

43 45 43 42 43 39 39 

42 43 

151 919 560 919 1225 1816 1711 1923 1793 2629 2838 

3.4 21 

12 21 

29 42 

49 68 63 


Number of Counties 

House finch count 
Index 2 

12 19 29 35 39 57 60 61 63 62 59 49 46 43 42 42 
5 9 30 109 298 316 625 934 1202 1280 1340 1398 1385 1763 2269 
0.26 0.31 0.86 2.8 5.2 5.3 10 15 19 22 27 30 32 42 54 

Christmas Index = House finch count/Number of Christmas counts. 
Index = House finch count/Number of counties reporting. 


Ecology: Weiss 

Vol. 106 (1997) 

2500 T 



o 1000 


-0 — Summer Count 
-B— Moving Average 


O r- CsJ 

00 CO 00 

91 O) O) 


Figure 5. Moving average plot of the Indiana summer house finch counts. 

count data and the count indexes demonstrate that the counts of wintering house 
finches in Indiana are growing at a greater rate than the May and summer counts. 
Inspection of the Christmas count data (Figure 1) and the Christmas count 
indices (Figure 2) reveals a curvilinear trend from 1980 to 1994 with a break 
occurring between 1988 and 1990. May counts show a similar trend with a break 
in 1989-1990. Summer counts show a sigmoidal growth pattern from 1980-1993, 
followed by a renewal of count increases in 1994. Moving average plots of the 
raw data from the summer and Christmas counts clearly show exponential growth 
through the 1980s that slowed in the early 1990s and then was followed by con- 
tinued growth into the mid-1990s (Figures 5 and 6). The May house finch data 
do not show an obvious slowdown in growth in the early 1990s (Figure 7). 


Ideally, small-bodied birds with a high reproductive potential and large brood 
sizes have an annual population growth potential of 50 to 100 percent (Ricklefs, 
1973). Counts of wintering house finches in Indiana are increasing faster than 

Vol. 106 (1997) Indiana Academy of Science 






| 3000 



o a- 

O t- <\l 

00 00 00 

CF> ff> ff> 











Figure 6. Moving average plot of the Indiana Christmas house finch counts. 

the May or summer counts; the winter counts are increasing exponentially with 
interruptions. This growth might represent the early stages of a sigmoidal growth 
curve for a population expanding into a new environment. House finch popula- 
tion growth east of the Mississippi based on Christmas count data from 1962 to 
1971 was exponential (Bock and Lepthein, 1976). Counts of individual house 
finches concentrated in wintering flocks would be expected to yield larger num- 
bers per unit effort than more dispersed populations that occur during either the 
spring migration or summer breeding season. Observers who watch bird feed- 
ers during the Christmas count should record most of the house finches tallied. 
The Indiana data, smoothed somewhat by using an index obtained by dividing 
the raw data by the number of counts (or participating counties), also revealed 
similar trends. 

Banding (Wiggins, 1987) and other studies (Hill, 1993) show that some house 
finches leave Michigan and other more northern localities during the winter. The 
southerly movement of the house finch into Indiana during the winter is 
reflected in the size of the Christmas counts. The wintering population is also 
increased by the finch's breeding success in the previous summer in Indiana and 
the surrounding States. May population counts consistently fall between the 
Christmas and summer counts, reflecting flock dispersal and the onset of the 
breeding season. Summer counts are consistently lower than either the Christ- 
mas or May counts. 


Ecology: Weiss 

Vol. 106 (1997) 

3000 t 






o a 

May Count 
Moving Average 






































Figure 7. Moving average plot of the Indiana May house finch counts. 

A number of density-dependent (recruitment, population size, dispersal, mor- 
tality, reproductive rate, competition, predation, and disease) and density-inde- 
pendent (climate, toxins, habitat alteration, backyard bird feeding, etc.) factors 
act to mediate growth and regulate house finch population size. The four prin- 
cipal ecological factors that limit bird populations are habitat, food supply, cli- 
mate, and disease (Gill, 1995). Subtle social forces, such as territory size, 
aggressiveness, dispersal rates, and recruitment, also influence the rate of pop- 
ulation growth. Recruitment of young birds into the local population varies 
inversely with adult mortality. The availability of winter feed to great tits (Van 
Balen, 1980) decreases the recruitment of young birds into the breeding popu- 
lation (lower adult mortality results in fewer vacancies in the local population), 
suggesting that backyard feeding may increase the dispersal rates of the house 

House sparrows, introduced into the U.S. in the early 1850s, benefitted from 
the existence of feed and grain stores as well as from horse droppings that con- 
tained undigested seeds. The coming of the automobile coupled with the decline 
in the number of feed stores stabilized house sparrow populations (Kastner, 1986). 
The recent increase in the popularity of backyard bird feeding has no doubt 
favored house sparrow populations as well as the populations of other species, 
including the house finch, that feed on the commonly served backyard bird foods. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 75 

In all seasons, 97% of the house finch diet is vegetable matter (Hill, 1993). 
The house finch prefers small sunflower seeds (oil) over milo, millet, or 
striped sunflower seeds, and, especially during winter months (Hill, 1993), ben- 
efits from the increasing popularity of backyard bird feeders (Bent, 1968). The 
use of feeders has stimulated a shift in the morphology of the house finch bill, 
allowing the finch to open sunflower seeds more efficiently (Sprenkle and Blem, 

The decline observed in the 1989 and 1990 Christmas counts may reflect 
the negative impact of the severe drought that persisted in the mid- to late 1980s 
on nesting success. The negative impact of the drought on the populations of 
other bird species was observed in Michigan and Wisconsin during the same 
time period (Blake, et ai, 1989). In addition, Bock and Lepthien (1976) found 
that the house finch does not prosper in extraordinarily wet years. The decline 
in the Indiana 1993 Christmas count may reflect the negative impact of the exces- 
sively wet 1992 breeding season on nest success. Eastern house finch popula- 
tions, however, seem to have adapted to wetter climates (Root, 1988). Christmas, 
May, and summer data for 1994 and summer data for 1995 indicate an increase 
in house finch numbers following the 1993 decline. 

House finches suffer from pox on their feet and legs, which, when spread to 
the bill and eyes, leads to blindness and death (Hill, 1993). A contagious con- 
junctivitis infection now being reported by bird banders in house finch popula- 
tions in the East may also lead to blindness and limit population growth. Competition 
with other bird species that occupy similar niches may also limit an otherwise 
explosive population increase in house finches. 


The regression equation for the Christmas counts projects a wintering house 
finch population of more than 10,000 birds by the year 2000. Continued expo- 
nential growth could take population size well above that figure. Continuation 
of the May trends would result in slightly more than 4,000 birds by the year 2000. 
and a continuation of the summer trends projects the presence of nearly 3.000 
birds by the year 2000. 

The ultimate impact of increased house finch numbers on other bird species 
may begin to reveal itself when house finch numbers reach levels closer to Indi- 
ana's carrying capacity for this and similar species. Are competitive interactions 
with house sparrows, American goldfinches, and other species that share simi- 
lar niche characteristics already limiting house finch population growth? What 
is the carrying capacity for the house finch in Indiana? How does backyard feed- 
ing effect the demographics of the house finch and other bird species? 

As with house sparrows in the 19 th Century, the arrival of house finches may 
be a mixed blessing. House finch depredations on commercial crops in Califor- 
nia and Hawaii have been documented (Hill, 1993) as has the negative impact 

76 Ecology: Weiss Vol. 106 (1997) 

of the house finch on house sparrow populations (Kricher, 1983). For bird enthu- 
siasts, the house finch endears itself through a more attractive, melodious song 
than the house sparrow. Are control measures for the house finch needed? 
More work is needed to determine the long-term effect of the house finch on 
other bird species as well as agricultural crops. 

The data collected by the dedicated volunteers who participate in the annu- 
al bird counts, although not always collected according to accepted scientific 
field methods, does provide a massive database for the many bird species that 
depend on Indiana habitats for their livelihood. Birds are sensitive indicators of 
the health of the environment, which justifies monitoring their population fluc- 
tuations and trends. A statewide program to standardize and train volunteers in 
data collection techniques would greatly enhance the value of future Christmas, 
May, and summer counts and provide the information to evaluate the status of 
Indiana's habitats well into the 21 st Century. 


The author would like to thank the hundreds of Indiana Audubon Society 
volunteers, compilers, editors, and publishers who have invested tens of thou- 
sands of hours over the years to gather and compile the data used for this 
analysis. The author would also like to thank the two anonymous reviewers who 
invested their time and effort to critique the manuscript. They provided encour- 
agement and many insightful suggestions for changes and improvements. Much 
of the scientific value of this paper is due to their valuable critiques. 


Blake, J.G., G.J. Niemi, and J.M. Hanowski. 1989. Drought and annual variation in bird populations. In: 

J.M. Hagan III and D.W. Johnson (Eds.), Ecology and Conservation of Neotropical Migrant Land- 
birds, pp. 419-430, Smithsonian Inst. Press, Washington, D.C., 609 pp. 
Bent, A.C. 1968. Life histories of North American cardinals, grosbeaks, buntings, towhees, finches, sparrows, 

and their allies. U.S. Nat. Mus. Bull. 237, 1,889 pp. 
Bock, C.E. and L.W. Lepthien. 1976. Growth in the eastern house finch population, 1962-1971. Amer. Birds 

30: 791-792. 
Clay, W.M. and K.M. Clay. 1981. A record of the house finch in southern Indiana. Indiana Audubon Quart. 

59(3): 91-92. 
Gill, EB. 1995. Ornithology, 2nd edition. W.H. Freeman and Co., New York, 766 pp. 
Hamilton, T.R. and CD. Wise. 1991. The effect of the increasing house finch population on house sparrows, 

American goldfinches, and purple finches in Indiana. Indiana Audubon Quart. 69(4): 251-254. 
Heller, D.D., Jr., and CD. Wise. 1982. The inevitable: House finch is found nesting in Indiana. Indiana Audubon 

Quart. 60(1): 6-13. 
Hill, G.E. 1993. House finch (Carpodacus mexicanus). In: A. Poole and E Gills (Eds.), The Birds of North 

America, No. 46, Acad. Natur. Sci., Philadelphia, and Amer. Ornithol. Union, Washington, D.C., 23 pp. 
Hopkins, E.M. 1980. 1980 May day bird count. Indiana Audubon Quart. 58(3): 84-96. 
Jackson, S.E 1983. 1981 summer bird count results. Indiana Audubon Quart. 61(1): 3-11. 
Kastner, J. 1986. A world of watchers. Alfred A. Knopf, New York, New York, 241pp. 
Kricher, J.C 1983. Correlation between house finch increase and house sparrow decline. Amer. Birds 37: 358- 

Mason, J. and A. Mason. 1982. The 1981 Christmas bird count. Indiana Audubon Quart. 60(2): 34-48. 
Ricklefs, R.E. 1973. Fecundity, mortality, and avian demography. In: D.S. Farner (Ed.), Breeding Biology of 

Birds, pp. 366-435, Nat. Acad. Sci., Washington, D.C, 515 pp. 

Vol. 106 (1997) Indiana Academy of Science 77 

Root, T. 1988. Atlas of wintering North American birds. An analysis of Christmas bird count data. Univ. Chica- 
go Press, Chicago, 312 pp. 

Sprenkle, J.M. and C.R. Blem. 1984. Metabolism and food selection of eastern house finches. Wilson Bull. 
96: 184-195. 

Van Balen, J.H. 1980. Population fluctuations of the great tit and feeding conditions in winter. Ardea 68: 

Wiggins, C.E. 1987. A plethora of house finches. Indiana Audubon Quart. 65(1): 37. 

Wise, CD. and A. Walls. 1988. A preliminary study of the impact of the house finch on populations of the 
house sparrow in Indiana. Indiana Audubon Quart. 66(1): 26-28. 

Proceedings of the Indiana Academy of Science 79 

(1997) Volume 106 p. 79-84 



John O. Whitaker, Jr., Rita McKenzie, Michelle Rakow, 

Brian Leibacher, and Priscilla Leibacher 

Department of Life Sciences 

Indiana State University 
Terre Haute, Indiana 47809 

ABSTRACT: Big brown bats first appeared in maternity colonies in mid- 
March and increased to peak adult populations by the first week of May. Exits 
began an average of 1.5 to 15.1 minutes after sunset. The length of time for 
all bats to exit averaged 18 minutes in the smallest colony (peak count of 69 
adults) to 44. 1 minutes in the largest colony (peak count of 349 bats). Tem- 
peratures below 10° C or rain caused bats not to emerge. The fall population 
decline started in late August and early September, but some bats remained in 
the colonies through mid-November. 

KEYWORDS: Big brown bat, Eptesicusfuscus, flight counts, populations. 


Most big brown bats (Eptesicusfuscus) hibernate alone in buildings in win- 
ter, at least in Indiana (Whitaker and Gummer, 1992; but also see Barbour and 
Davis, 1969 and Baker, 1983). Some hibernate in caves and mines (Beer, 1955; 
Beer and Richards, 1956; Phillips, 1966). The bats often move within and between 
roosts during the hibernation period (Goehring, 1972; Mills, et al., 1975; Whelden, 
1941). They start to form maternity colonies in March, and the colonies disband 
by early November (Mills, et ah, 1975; Whitaker and Gummer, 1992). How- 
ever, few specific details on the buildup and decline are available. 

The objectives of the present study were to examine changes in the number 
of big brown bats emerging at dusk from maternity colonies as related to sea- 
son, weather, and bat biology. Specifically, we wanted to determine: (1) the dates 
of the beginning, progression, and completion of the spring increase and of the 
abandonment in the fall at maternity colonies; and (2) fluctuations in flight counts 
due to weather, the young becoming volant, bats moving, or other causes. 


Observations were made by Brian and Priscilla Leibacher at the Scotland 
Hotel in Greene County; by Darrel and Rita McKenzie at the Presbyterian Church 
in Roachdale, Putnam County (and two observations at the Jamestown church): 
and by Michelle Rakow at Mecca School in Parke County. The exits from the 
roosts were watched, and all the exiting bats were counted from approximately 
{ h hour prior to sunset until 15 minutes after the last bat left the roost. The data 

80 Ecology: Whitaker, et al Vol. 106 (1997) 

recorded included the exit time of the first bat, the number of bats exiting each 
minute, temperature, wind, cloud cover, and the time the last bat exited. Tem- 
perature data were obtained from the National Weather Service and from on-site 
readings. All data were collected in 1989, except for supplementary data col- 
lected at the Scotland Hotel in 1995. 


Jamestown and Roachdale. These colonies were both in churches. Only 
two observations were made at Jamestown (Table 1): the first was on 17 March, 
when 20 bats emerged, and the second was on 26 March, when 40 bats emerged. 
The first observations at Roachdale were on 26 March (23° C), when 24 bats 
emerged. None were seen on 7 April, probably because of the low temperature 
(6° C). On 14 April (13° C), 50 emerged; on 25 April (22° C), 148 emerged; and 
on 4 May (14° C, rain), 58 were counted. Maximum counts at Roachdale (Table 
1) were 349 bats on 14 August (with young), and 306 bats on 14 May (pre-young). 
The Roachdale site is also used for hibernation by low numbers of bats (up to 5 
per winter). 

Scotland Hotel. This colony was in the attic of an old hotel. Observations 
were made on 15 March (temperature 4° C, cloudy), 23 March (11° C, clear), 
30 March ( 1 1 ° C), 5 April ( 1 0° C, drizzle), 1 2 April ( 1 1 ° C, drizzle), and 20 April 
(17° C), but no bats were seen until 20 April, when 45 exited. The temperature 
and weather conditions on both 23 and 30 March were such that bats would have 
exited if present (insects were flying). The bats apparently had not begun return- 
ing by 30 March. No bats were seen on 5 or 12 April; however, it was raining on 
both those nights, which might have kept the bats from exiting. Unfortunately, 
no other nights were sampled. Therefore, we can only say that the first bats appar- 
ently returned between 30 March and 20 April. No bats hibernate at the hotel, 
as it is not heated in winter. 

In 1989, 26 bats emerged on 12 October (15° C), 17 on 25 October (19° 
C), and none on 8 November (11° C). The bats had apparently left by the last 
date. Supplementary data were collected in 1995 (Table 1). Seventy-two bats 
exited on 4 September, but this number rapidly declined. The last bats seen exit- 
ing numbered 2 on 23 October (12° C). None exited on three nights in Novem- 
ber or on 1 December. In 1995, bats were seen exiting from a heated building 
near the Scotland Hotel. The number exiting that building was 34 and 47 in 
late September, from 5 to 9 between 10 and 18 October, and 44 on 18 October. 
On 23 October and 2 November, 8 and 3 bats exited this building; none were 
observed on 9 and 13 November (temperatures 3° C and 4° C, respectively). 
However, on 1 December, 10 bats exited the building. Big brown bats exit in 
winter, especially on warm nights, and we believe these latter bats had begun 

Mecca. The colony at Mecca was in the attic of an old, but refurbished, 
school. Observations at Mecca School paralleled those at the Scotland Hotel. No 
bats were seen there during observations on 16 March (temperature -5° to 13° 

Vol. 106 (1997) Indiana Academy of Science 


Table 1. Eptesicus emergence counts at the Scotland (* = 1995 data), Mecca, and Roach- 
dale (2 at Jamestown = **) colonies. Temperatures are in centigrade. R or C after the 
date indicates Rain or Cloudy. 




remp No. 

Exit times 

Temp No. 

Exit times 



Exit times 



















































26 R 








































22 R 












29 C 























26 C 





























31 C 
































27 C 































25 C 















































24 C 




























12 C 













82 Ecology: Whitaker, et al. Vol. 106 (1997) 

C, min/max Rockville), 23 March (-2° to 14° C), 31 March (2° to 10° C), and 7 
April (0° to 13° C); the first bats were seen on 29 April (14° to 23° C), when 195 
emerged. The bats returned between 7 and 29 April. The maximum number of 
bats prior to flight by the young was 242 on 5 May; this was also the overall 
maximum (Table 1). About 6 to 20 bats regularly hibernate at Mecca. 


Establishment of Maternity Colonies in Spring. Bats began returning to 
two of the maternity colonies by at least 17 March (Jamestown) and 24 March 
(Roachdale), whereas limited data indicate the arrival of the first bats at Scot- 
land between 30 March and 20 April. The fact that 45 (more than half the ulti- 
mate total) emerged on 20 April suggests that the first bats probably arrived at 
the colony in Scotland closer to 30 March than to 20 April. At Mecca, the spring 
buildup began between 7 April and 29 April. Their arrival was probably much 
closer to 7 April as the full complement was there on 29 April. Big brown bats 
start the spring buildup in central Indiana in March or early April. In the 
spring, the full complement of bats was attained by about 14 May at Roachdale 
and 3 May at Scotland and Mecca. 

Exit Times. In 1989, exit time showed a pronounced correlation with sun- 
set time. The bats usually emerged between 1 and 15 minutes after sunset (ear- 
lier on overcast days). Exceptions occurred on 4 May at Roachdale, when the 
first bat exited at 7:30 RM. (sunset at 7:47; misty; and the count was low), and 
on 5 July, 7 minutes before sunset (no weather information). However, at the 
Scotland Hotel on the same day (5 July), the first bat exited 6 minutes after 
sunset. The first bats leaving the hotel on that date flew erratically and may have 
been young. 

Amount of Time to Exit. During the 18 nights on which more than 1 bat 
exited in Roachdale, the number emerging ranged from 39 to 316 (# = 147.4). 
The bats took from 22 to 45 minutes to emerge, averaging 35.3 minutes. No bats 
emerged on 7 April, when the temperature was 6° C, and only one emerged on 
22 May, when it was raining. The number exiting the Scotland Hotel averaged 
44.8 (8-91 bats, n = 16) in 1989. The bats took an average of 17.9 minutes to exit 
(ranging from 8 to 37 minutes). On the one night when it rained at exit time, 
none emerged. The bats averaged 16.3 minutes to emerge in the autumn of 1995. 
On 14 nights, the number emerging from the Mecca School ranged from 7 to 
242 (% = 170.1). The bats averaged 25.4 minutes to exit (ranging from 3 to 49 
minutes). On 4 October, the temperature was 3° C, and only 3 bats exited (exit 
time 3 minutes); on 18 October, the temperature was 4° C, and none exited. How- 
ever, a week later, 66 exited when the temperature was 21° C. Cold, rainy nights 
clearly deter bats from exiting. 

Temperature. Exceptions exist, but temperature clearly influences the num- 
ber of bats exiting. For example (Table 1), at Roachdale, 25 bats were counted 
on March 26, when the temperature was 23° C, but one week later with a tem- 
perature of 6° C, no bats exited. Lower temperatures on which bats emerged 

Vol. 106 (1997) Indiana Academy of Science 83 

were 3° C on 4 October at Mecca, when seven emerged; 14° C at the Scotland 
Hotel on 10 May, when 54 emerged; and temperatures of 14°, 9°, 12°, and 13° 
C at Roachdale when 58 (4 May), 55 (6 October), 39 (13 September), and 1 
(22 May, a rainy night) emerged. Large numbers of bats generally did not emerge 
at temperatures much below 10° C. Usually, fewer bats exited at temperatures 
less than 15° C, except for 14 May at Roachdale, when over 300 emerged at a 
temperature of 13° C. The same was true at Mecca, where on 10 May approxi- 
mately 240 bats were counted when the temperature was 13° C. On 4 October, 
also at Mecca (3° C), seven exited; on 18 October (4° C), none exited; on 25 
October (21° C), 66 were counted. On 1 November (3° C), bats were heard, but 
they did not exit. The temperatures were somewhat low (ranging from 9° to 
15° C) at Roachdale between 13 September and 16 October, and the number of 
bats emerging was correspondingly lower, ranging from 39 to 72. However, on 
24 October, 118 bats emerged at a temperature of 20° C. The Scotland Hotel 
seemed to have an erratic count relative to temperature. After the first exit count 
of 45 on 20 April (17° C), no bats emerged on 26 April, when it rained; high 
counts occurred on 3 May (15° C; 69 bats) and 10 May (14° C; 54 bats). How- 
ever, no more than 15 bats emerged on 17 May, 24 May, and 2 June, even though 
the temperature was above 21° C, and there was no rain. Perhaps many of the 
bats were at an alternate location at that time. No bats exited from the Scotland 
Hotel or the nearby building on 1 or 8 November, 1989, when the temperature 
was 11° C. However, bats did emerge from the nearby building on 1 December 
at a temperature of 10° C. 

Rain. It rained on 5, 12, and 26 April at the Scotland Hotel (Table 1), and 
no bats emerged. Since no bats had been seen at this colony by 12 April, they 
might not have been present. Forty-five bats emerged on 20 April at the Scot- 
land Hotel, but none emerged on the night of 26 April, when it rained. On suc- 
cessive dates at Roachdale (25 April, 4 May, 14 May, 22 May, and 29 May), 
the bats emerging numbered 148, 58, 306, 1, and 176, respectively. It rained on 
4 and 22 May, the two nights when numbers were greatly reduced. Rain clear- 
ly reduced the number of bats emerging. Sunset apparently has a greater influ- 
ence on exit time than temperature, although very low temperatures or rain might 
deter the bats from exiting; cloudy conditions may cause early departures. 

Fall Decline. The bat populations started declining in August or September. 
By 2 August, the number leaving the Scotland Hotel was reduced to 58 and then 
tended to level out with 55 being seen on 16 August and 54 on 3 1 August. On 7 
September, the exit count was only 5; on 12 October, the number was 26; on 17 
October, 25; and no bats emerged on 8 November, even though the tempera- 
ture was 10° C. The major drop in count at Mecca from 122 on 16 August (23° 
C) to 29 on 13 September (15° C) followed by an increase to 212 on 20 Sep- 
tember (19° C) is hard to understand, as the temperature was relatively high on 
all three nights. No bats emerged at Mecca on 1 and 8 November (3° and 10° 
C, respectively). The Roachdale Church bats arrived earlier (26 March) than 
those at the other two main sites, although the earliest arrival date was at Jamestown. 

84 Ecology: Whitaker, et al Vol. 106 (1997) 

The bats also left Roachdale later (12 November count of 53; 10° C) and may 
have been there even later. Bat numbers declined, and most of the bats had left 
the maternity colonies by early November. 


Baker, R.H. 1983. Michigan mammals. Michigan State Univ. Press, East Lansing, 642 pp. 

Barbour, R.W. and W.H. Davis. 1969. Bats of America. Univ. Press Kentucky, Lexington, 286 pp. 

Beer, J.R. 1955. Survival and movements of banded big brown bats. J. Mammal. 36: 242-248. 

and A.G. Richards. 1956. Hibernation of the big brown bat. J. Mammal. 37: 31-41. 

Goehring, H.H. 1972. Twenty-year study of Eptesicus fuscus in Minnesota. J. Mammal. 53: 201-207. 

Mills, R.S., G.W. Barrett, and M.P Farrell. 1975. Population dynamics of the big brown bat {Eptesicus fus- 
cus) in southwestern Ohio. J. Mammal. 56: 591-604. 

Phillips, G.L. 1966. Ecology of the big brown bat (Chiroptera: Vespertilionidae) in northeastern Kansas. Amer. 
Midi. Natur. 75: 168-198. 

Whelden, R.M. 1941. Hibernation of Eptesicus fuscus in a New Hampshire building. J. Mammal. 22: 203. 

Whitaker, J.O., Jr., and S.L. Gummer. 1992. Hibernation of the big brown bat, Eptesicus fuscus, in buildings. 
J. Mammal. 73:312-316. 

Proceedings of the Indiana Academy of Science 85 

(1997) Volume 106 p. 85-94 





Raymond A. Cloyd, C. Richard Edwards, and Larry W. Bledsoe 

Purdue University 

1158 Entomology Hall 

West Lafayette, Indiana 47907-1158 

ABSTRACT: The pod trichome densities of eight soybean lines and one pub- 
lic variety were determined, and the associated aspects of bean leaf beetle, 
Certoma trifurcata (Forster), feeding behavior on soybean pods were inves- 
tigated. Three-seeded soybean pods were removed from the top and the bot- 
tom of the plants of selected lines. The pods were divided into seven sections, 
starting from the peduncle and ending at the pod tip. Circular areas within 
each section were delineated, and the number of trichomes in each circle 
was determined. Data were analyzed for differences in trichome numbers 
among lines, within plant regions, and among sections of pods. The tri- 
chome density for different lines ranged from 495 to 924 trichomes/cm 2 . Pods 
from the top of the plant had higher trichome densities than pods from the bot- 
tom. Soybean pod sections nearest the pod tip had the highest trichome den- 
sities, and the sections nearest the peduncle had the lowest. Free choice feeding 
assays were conducted in the laboratory with three-seeded pods from all lines. 
Field-collected adult bean leaf beetles were allowed to feed for forty-eight 
hours. The percentage of the surface damaged per pod section was estimated. 
Bean leaf beetles showed a stronger tendency to feed on pods at the pedun- 
cle end and the sections nearest it. These pod sections had significantly lower 
numbers of trichomes. Areas with the highest trichome densities had the low- 
est percentage of beetle damage. Lines HC83- 193-5 and HC83- 123-9 and the 
experimental cultivar "Anderson Velvet" had the greatest resistance to over- 
all pod damage. 

KEYWORDS: Bean leaf beetle, feeding behavior, host-plant resistance, pods, 
pubescence, soybeans, trichomes. 


Soybean, Glycine max (L.) Merrill, is attacked by many foliar and pod feed- 
ing insects (Kogan, et ai, 1988). The bean leaf beetle, Cerotoma trifurcata 
(Forster) (Coleoptera: Chrysomelidae), causes pod injury that predisposes the 
seeds to injury by secondary pathogens (Shortt, et ai, 1982) such as Alternaria 
tenuissima (Kunze ex Pers.), resulting in yield and seed quality reductions (Smelser 
and Pedigo, 1992a). The feeding behavior of first and second generation beetles 
is different. Second generation adults prefer to feed on soybean pods instead of 
leaves (Sims, et ai, 1984). The larvae feed on roots, root hairs, and Rhizobium 
nodules (McConnell, 1915). Bean leaf beetles are also important vectors of bean 
pod mottle virus in the southern United States (Hopkins and Mueller. 1983). 

86 Entomology: Cloyd, Edwards, and Bledsoe Vol. 106 (1997) 

Bean leaf beetle management relies mainly on the use of insecticides. Despite 
their effectiveness, insecticides can create environmental problems. Research on 
alternative management strategies that minimize insecticide application has 
focused on the use of resistant plant material. This approach also has the poten- 
tial to delay the development of insecticide resistance in the bean leaf beetle. Tri- 
chomes on foliage and/or pods can act as a physical barrier to inhibit insect 
feeding (Norris and Kogan, 1980). 

Trichome density, the number of trichomes per unit area, has been shown to 
influence the behavior of several chrysomelid beetles. Lamb (1980) found that 
the trichomes on the pods of the mustard plant inhibited feeding by the flea bee- 
tle, Phyllotreta cruciferae (Goeze). Flea beetle feeding increased on pods whose 
trichomes were removed. Baur, et al. (1991) demonstrated through the use of 
dual-choice laboratory assays that trichome density influenced Agelastica alni 
L. oviposition and feeding on the gray alder, Alnus incana (L.). Palaniswamy 
and Bodnaryk (1994) showed that wild Brassica species having a leaf tri- 
chome density greater than 2,172 trichomes/cm 2 were highly resistant to flea 
beetle feeding. Brassica species with less than 30 trichomes/cm 2 suffered sig- 
nificant damage. Behavioral observations showed that high trichome densities 
act as a physical barrier to flea beetle feeding. A notable case of the influence 
of trichome density on a chrysomelid beetle is the effect of wheat leaf trichome 
density on the behavior of the cereal leaf beetle, Oulema melanopus (L.). Tri- 
chome density influenced cereal leaf beetle oviposition (Gallun, et al., 1966; 
Casagrande and Haynes, 1976; Lampert, et al, 1983), egg viability (Lampert, 
et al, 1983), and larval survival (Ringland and Everson, 1968; Hoxie, et al, 
1975). Schillinger and Gallun (1968) found that pubescence deterred adult cere- 
al leaf beetle oviposition on wheat leaves and resulted in abnormal behavior {e.g., 
movement). Fewer eggs hatched on highly pubescent leaves, suggesting that the 
eggs may have become desiccated. In addition, trichome density reduced early 
larval stage survival. The larvae have to eat through the trichomes to reach the 
leaf surface. The composition of the trichomes ingested {e.g., lignin and cellu- 
lose) caused an imbalance in the larval diet (Schillinger and Gallun, 1968). 

Minimal information exists on trichome density and its effects on soybean 
pod feeders. The major objectives of this study were to (1) determine if differ- 
ences exist in soybean pod trichome density and to (2) examine any relationship 
between trichome density on soybean pods and feeding behavior by adult bean 
leaf beetles. 


Trichome Density. Eight soybean lines (MBB80-169, MBB83-190, MBB80- 
133, MBB83-368, HC83-193-5, HC83-19-2, HC83-123-9, and L76-0038) known 
to demonstrate foliar resistance to the Mexican bean beetle, Epilachna varivestis 
(Mulsant) and one susceptible cultivar, "Williams 82," were planted on 28 
May 1993 at the Purdue University Agronomy Research Center, West Lafayette, 
Indiana. Each line and cultivar was sampled and analyzed using a completely 

Vol. 106 (1997) Indiana Academy of Science 


Figure 1 . A diagram of a three-seeded soybean pod showing the areas where trichomes 
were counted. 

randomized experimental design. The experimental area was 222.8 m 2 (12.1 m 
x 18.2 m) and previously had been planted to corn. Each line was planted in two- 
row plots 154.9 cm long and 76.2 cm wide, oriented north to south. Plots were 
separated by a 30.4 cm east to west alley. Blocks were separated on the ends with 
alleys 121.9 cm wide. Sixty-four seeds of each entry were planted per row with 
a Planet Junior® Sample Plot Planter (Swanson Machine Co., Champaign, Illi- 
nois) at a depth of 0.63 to 1.2 cm. Plants of "Anderson Velvet" were planted in 
a single row by themselves in the same area. 

On 24 June 1993, a 18 x 14 mesh polyethylene screen (Lumite Co., Gainesville, 
Georgia 30503) was placed on a metal frame over the entire area to prevent 
insects such as the Japanese beetle (Popillia japonic a Newman), bean leaf bee- 
tle, and grasshoppers (various species) from feeding on the foliage. 

On 9 September 1993, approximately 30 pods were randomly collected from 
the top and 30 from the bottom of 5 randomly selected plants in the R6 growth 
stage (Ritchie, et ah, 1992). Pods were collected and placed into polyethylene 
bags (15.2 x 7.6 cm), transported to the laboratory, and stored at 2.8 ± 2° C. 

Laboratory Evaluation. In the laboratory, 5 three- seeded pods per line were 
randomly selected from the 30 pods collected from the top and bottom of the 
plants from each line. Pods were divided into seven sections starting at the pedun- 
cle and ending at the apex of the pod (Figure 1). The section between the pedun- 
cle and the first seed was represented by the letter "A." Each of the three sections 
of the pod containing a seed were divided into an upper and lower section (near- 
est the dorsal and ventral sutures) represented by the letters "B" and "C," respec- 
tively. Circular areas within each section were delineated with a 0.5 cm cork 
borer (0.2 cm 2 ). The number of trichomes in each circle was counted with the 
aid of a 20x dissecting microscope. Data were analyzed by ANOVA (SAS 
Institute, 1988). The Student-Newman-Keuls Sequential Range Test (SNKS 
Range Test) was used to separate significant differences {P < 0.05) among lines, 
plant regions, and pod sections. 


Entomology: Cloyd, Edwards, and Bledsoe Vol. 106 (1997) 

Table 1 . The mean pod trichome density among selected 
soybean lines. Within column means followed by the same 
letter do not differ significantly (Student-Newman-Keuls 
test: P < 0.05, n = 70). 


No. of trichomes / cm 2 


924.3 a 


831.7 ab 


803.7 a b 


794.0 a b 


747.0 b 


669.0 b c 


598.2 c d 


548.5 c d 

Williams 82 

495.2 d 

Bean Leaf Beetle 
Feeding Behavior 
Assays, 1993. Adult 
feeding preference 
assays were conduct- 
ed on 9, 18, and 27 Sep- 
tember and 7 October. 
Pods used for the assay 
were collected within 
24 hours of the assay. 
Thirty three-seeded pods 
were collected from the 
top and 30 from the bot- 
tom of each line. Three- 
seeded pods of 
approximately the same size from seven of the lines plus "Anderson Velvet" 
were randomly placed on filter paper moistened with distilled water. The pods 
were placed equidistant around the perimeter of a Petri dish (2.5 x 15.2 cm). Fif- 
teen field-collected adult bean leaf beetles of unknown sex and age were 
placed in each Petri dish and allowed to feed for 48 hours. The Petri dishes were 
placed on a laboratory bench at 22.2 ± 5° C at a 14:10 (L:D) photoperiod with 
a relative humidity of 85 ±5% inside the Petri dishes. Three Petri dishes, each 
representing a replication, were evaluated each day. 

Each pod was divided into 7 sections (see above). Both surfaces of the pod 
were evaluated. Feeding damage was visually estimated as the percentage of pod 
surface damaged per pod section. Average damage per section was determined 
by dividing the mean values by two, since both surfaces of the soybean pod were 
evaluated for damage. Average damage per pod was determined by summing the 
means of the percent damage per section for each line and dividing by the 
number of sections (n = 7). Pod section means for each line were determined 
by summing all the percent damage per section means for each section and divid- 
ing this value by the number of lines (n = 8). Due to the irregularity and round- 
ness of the soybean pod, the actual area damaged by the bean leaf beetles was 
not measured. Because the data from the feeding assay studies were based on 
subjective visual observations on non-standard experimental units, formal sta- 
tistics were not applied. However, the results were consistent for both years of 
the study, and a distinct range of feeding by the adult bean leaf beetles was noted 
in all assays (Tables 3-5). This information suggests that a feeding pattern exists. 
Bean Leaf Beetle Feeding Behavior Assays, 1994. Adult feeding prefer- 
ence assays were conducted on 9, 12, 14, and 21 September. Pods used for this 
study were collected within 24 hours of the assay from the Purdue University 
Agronomy Research Center and the Environmental Entomology Laboratory. 
Identical lines were included in each assay. The experimental cultivar "Ander- 
son Velvet" and the glabrous line D88-5320 were also included in the 1994 assay. 
The bioassay method was identical to that described above. 

Table 2. The mean 

trichome density of pod sections aver- 

aged over soybean 

lines/cultivars. Within column means 

followed by the same letter do not differ significantly (Stu- 

dent-Newman-Keuls test: 

P < 0.05, n = 90). 


No. of trichomes / cm 2 


971.0 a 


831.0 b 


811.9 b 


671.8 c 


650.5 c 


544.8 d 


505.6 e 

Vol. 106 (1997) Indiana Academy of Science 89 


Trichome Densi- 
ty Study. Significant 
differences (P < 0.05) 
occurred in trichome 
densities among lines 
(F= 12.0, df= 8, 72), 
plant regions (F = 7.2, 
df - 1, 72), and pod 
sections (F = 365.9, 
df= 6,432). HC83-193- 
5 had the highest tri- 
chome density, while 
"Williams 82" had the lowest (Table 1). Significantly (P < 0.05) higher trichome 
densities were found on the pods at the top of the plant (759.8 trichomes/cm 2 ) 
than at the bottom (684.8 trichomes/cm 2 ). Trichome densities also varied sig- 
nificantly (P < 0.05) among sections of the soybean pod (Table 2). Section 3B 
had a significantly higher trichome density than other sections, followed by sec- 
tions 2B and 3C, 2C and IB, 1A, and 1C. Section 1C had a significantly lower 
trichome density than all other sections. 

Differences in trichome density among lines may be the result of genetic 
variation. Bernard and Weiss (1973) reported that genetically controlled varia- 
tion in leaf trichome density occurs in some exotic soybean germplasm. Zaiter, 
et al. (1990) demonstrated that trichome density is under genetic control and that 
differences in leaf trichome densities exist among soybean cultivars. 

Differences in trichome density among plant regions is not unusual. Yapp 
(1912), working with Spiraea ulmaria L., and Stober (1917), working with herbs, 
reported that leaves on the upper part of the plant have more trichomes than 
leaves located toward the base. Ehleringer and Mooney (1978) reported that leaf 
pubescence reduces light absorptance and lowers heat loads. Reflection of solar 
radiation maintains the leaf temperature below the air temperature resulting in 
lower transpiration rates. Because the upper region of the plant receives higher 
amounts of direct solar radiation, trichomes may act as a barrier to reduce the 
amount of light reaching the pod surface, thus reducing moisture loss and des- 
iccation. In addition, as demonstrated with soybean leaves, trichomes reduce 
wind movement which lowers transpiration rates from leaf surfaces (Woolley, 
1964). Pods in the top region of the soybean plant are exposed to wind, thus 
increased trichome density would presumably reduce water loss. The plant canopy 
reduces wind movement among pods at the bottom of the plant creating a micro- 
environment in which trichome density is less important for reducing transpi- 
ration from the pod. 

The differences in the trichome density on many plants could result from 
surface area expansion. Some young organs, such as leaves, fruits, and stems, 


Entomology: Cloyd, Edwards, and Bledsoe Vol. 106 (1997) 

Table 3. The percent adult bean leaf beetle damage per pod section corresponding to 
1993 and 1994 assay dates. 






9/12/94 9/14/94 







25.9 32.8 







12.9 18.0 







15.7 18.7 







6.0 4.6 







4.4 5.7 







4.8 5.0 







9.8 8.5 


Table 4. The percent adult bean leaf beetle damage per 

pod (= line) corresponding to 

1993 assay dates 







Williams 82 












Anderson Velvet 














Williams 82 


Williams 82 
















Anderson Velvet 


Anderson Velvet 


have a dense covering of trichomes. As the organ grows, the trichomes become 
spaced further apart, and, if new trichomes are not produced, growth results in 
lowered trichome densities (Johnson, 1975). With soybean, the continued expan- 
sion of the pod surface (the number of cells increases, and these cells elongate) 
moves the trichomes further apart and reduces trichome density (Jackai and Oghi- 
akhe, 1989). At the time of sampling, the pod sections nearest the peduncle (1 A, 
IB, and 1C) had undergone more expansion than those farther from these sec- 
tions, resulting in the gradient of trichome density over the length of the pod. 
Because expansion was less at the apex of the pod, sections closer to the apex 
(2B, 2C, 3B, and 3C) had higher trichome densities. 

Bean Leaf Beetle Feeding Behavior Studies. Adult bean leaf beetles demon- 
strated a strong preference for feeding at the pod's peduncle end and the sections 
nearest it (Table 3, columns 2-4 and 5-8). The greatest damage occurred in sec- 
tions 1 A, 1C, and IB, which had the lowest trichome densities (Table 2). The 
least damage occurred in sections 2B and 3B, which had the highest trichome 
densities. Field observations have shown that adult bean leaf beetles tend to feed 
on the peduncle end of soybean pods (L. Bledsoe, pers. comm.). The results of 

Vol. 106 (1997) Indiana Academy of Science 91 

Table 5. The percent adult bean leaf beetle damage per pod (= line) corresponding to 
1994 assay dates. 






Percent Damage 
Per Pod 

(Ave. /Year) 

Williams 82 










































Anderson Velvet 
























these laboratory studies suggest that bean leaf beetles feed on the peduncle end 
and the sections nearest it because these sections have the lowest trichome 
densities. In addition, bean leaf beetles may be able to "mow" down the trichomes 
in these sections, making it easier to reach the pod's outer surface. Hulley (1988) 
demonstrated this behavior with the caterpillar, Pardasena sp. nr diver sip ennis 
Gaede, on Solarium coccineum Jacq. 

Lines HC83-123-9 and HC83-193-5 as well as 'Anderson Velvet" demon- 
strated the greatest resistance to adult bean leaf beetle pod feeding (Table 4 and 
5). "Anderson Velvet" is highly pubescent. The high number of trichomes made 
it difficult to obtain reliable counts, so trichome density was not determined. The 
unsuitability of the various lines to bean leaf beetle feeding might also result 
from nutritional deficiencies or disproportionalities, biophysical deterrence, or 
physiological inhibitors (Barney and Rock, 1975). In addition, other genetical- 
ly controlled factors might influence the degree of resistance (Clark, et «/., 1972). 

Although more studies have been conducted to investigate the interaction 
of soybean leaf trichome density than pod trichome density with insect resis- 
tance (Chiang and Norris, 1983; Zhan, et al., 1986; Gunasinghe, et al.. 1988; 
Lambert, et ah, 1992), research with other agricultural crops has demonstrat- 
ed that pod trichome density is associated with reduced feeding by some insects. 
Chiang and Singh (1988) found that trichomes on the pods of cowpea. Vigna 
vexillata, contributed to resistance against the pod-sucking bug, Clavigralla 
tomentosicollis Stal. Jackai and Oghiakhe (1989) reported that trichomes on the 
pods of two cowpea varieties interfered with pod feeding and development of 

92 Entomology: Cloyd, Edwards, and Bledsoe Vol. 106 (1997) 

the legume pod-borer, Maruca testulalis (Geyer), as well as C. tomentosicollis. 
They found that the insects were restless and had difficulty positioning their 
legs on the pod wall. Oghiakhe, et al. (1992) determined that greater trichome 
density on Vigna unguiculata pods reduced larval damage from M. testulalis 
(Geyer), and they concluded that trichome density was the major factor con- 
trolling resistance. 

While some studies have demonstrated the effectiveness of soybean pod tri- 
chomes in reducing insect feeding damage, other studies show that increased 
pubescence is not always associated with resistance. Morse and Cartter (1937) 
found that glabrous soybeans were highly resistant to the soybean pod borer, 
Laspeyresia glycinivorella Mats., whereas pubescent varieties were highly sus- 
ceptible. Broersma, et al (1972) reported that soybean strains with 2,969 to 3,100 
trichomes/cm 2 had significantly higher potato leafhopper populations than strains 
with 610 to 810 trichomes/cm 2 . Turnipseed (1977) observed that potato leafhop- 
per populations were less affected by trichome density than trichome length. 

Very little information was found in the literature on the interaction of pod 
trichome density and bean leaf beetle feeding damage. Previous studies have 
only investigated the effect of pod pubescence on reducing seed quality loss due 
to bean leaf beetle feeding and subsequent invasion by Alternaria tenuissima 
(Shortt, et a/., 1982) and in relation to yield and quality reduction (Smelser and 
Pedigo, 1992a, b). 

The existence of an inverse relationship between increased trichome densi- 
ty and feeding preference by bean leaf beetles for both soybean lines (Tables 4 
and 5) and pod sections within the lines (Table 3) was found in this study. HC83- 
123-9 was a notable exception to this trend. This line showed a high level of 
resistance to bean leaf beetle feeding at a much lower pod trichome density (Table 
1). HC83- 123-9 is highly resistant to Mexican bean beetle and adult Japanese 
beetle foliar feeding (Cooper and Hammond, 1988), suggesting that factors other 
than trichome density may contribute to host plant resistance. Nevertheless, high 
trichome densities may inhibit adult bean leaf beetles from reaching the pod sur- 
face and feeding. Additional studies should be conducted to evaluate this behav- 
ior further. 


The authors gratefully acknowledge Dr. Roger H. Ratcliffe, Dr. Clifford S. 
Sadof, Dr. Richard E. Shade, and Dr. Alan C. York for their comments on the 


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agromyzid beanflies. Environ. Entomol. 12: 260-265. 

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Proceedings of the Indiana Academy of Science 95 

(1997) Volume 106 p. 95-104 




J. Pichtel, Angela Covey, and K. Lukscay 

Department of Natural Resources and Environmental Management 

Ball State University 

Muncie, Indiana 47306-0495 

ABSTRACT: Ethylenediaminetetraacetic acid (EDTA) and nitrilotriacetic 
acid (NTA) were evaluated in column studies (each 0.1, 0.01, or 0.001 M) for 
their ability to extract lead (Pb) and chromium (Cr) from contaminated soil 
(Pb tot = 1300 mg/kg; Cr tot = 4940 mg/kg; pH = 10.3) collected at an abandoned 
industrial facility. The EDTA was eluted at pH 3.0, 5.5 (ambient), and 10.0, 
and the NTA at pH 3.0 and 11.1 (ambient). The efficiency of Pb and Cr solu- 
bilization was influenced by solution pH and chelant-metal chemistry; the 
EDTA, a hexadentate ligand, solubilized both metals more effectively than 
did the quadridentate NTA. Lead and Cr removal increased with higher EDTA 
concentrations; however, higher NTA concentrations did not remove signifi- 
cantly greater amounts of Pb or Cr (P < 0.05). The 0. 1 M EDTA at pH 3.0 and 
10.0 removed 86.5% and 87. 1% Pb after 200 pore volumes, respectively. The 
0.1 M NTA (pH 3) recovered 29.5% soil Pb. The 0.1 M EDTA, pH 5.5, recov- 
ered 40.9% soil Cr, and the 0. 1 M NTA at pH 1 1 . 1 removed 28.5%. The lower 
Cr removal compared to that for Pb may be explained by the chemical forms 
of each metal in the soil: 89.8% Cr occurs in residual forms, which are not 
readily extractable except by exhaustive processes. In contrast, only 45.7% 
Pb occurs in residual form. Initial flushing with 0.1 N HC1 followed by 
chelant flushing did not significantly improve removal of either Pb or Cr. Pb 
removal via soil flushing was equal or greater in effectiveness to that by ex- 
situ processes. 

KEYWORDS: Chromium, EDTA, lead, NTA, soil flushing. 


Soils at numerous industrial sites are contaminated with heavy metals 
(Richards, et a/., 1993; Cairney, 1987). Metal-rich sites pose potential hazards 
to public health and the environment via contamination of groundwater and sur- 
face water as well as through plant uptake. 

The use of technologies which eliminate or reduce the hazardous charac- 
teristics of waste is now being given greater priority over traditional contami- 
nant removal methods, such as excavation followed by landfilling. Available 
treatment technologies for remediating metal-contaminated soils include solid- 
ification/immobilization processes, soil washing {ex-situ), and soil flushing 

Metals on weathered metalliferous sites occur in complex forms (Kabata- 
Pendias and Pendias, 1992), and their mobility is controlled by several chemi- 
cal and physical phenomena, including soil pH, soil type, cation exchange capacity. 

96 Environmental Quality: Pichtel, et al. Vol. 106 (1997) 

particle size, contaminant concentration, and the presence of organic and inor- 
ganic compounds. Many of these factors are interdependent (Reed, etal, 1995). 
Metal removal efficiencies during soil flushing depend not only on soil charac- 
teristics but also on metal characteristics, extractant chemistry, and processing 
conditions. Chelating agents, such as ethylenediaminetetraacetic acid (EDTA) 
and nitrilotriacetic acid (NTA), bond with the metal to facilitate solubilization 
in the extraction medium. The ability to form stable metal complexes makes 
EDTA and NTA effective extractants for metal-contaminated soils (Davis and 
Singh, 1995; Elliott and Brown, 1989; Cline, et al., 1993). 

Chelants vary in effectiveness for Pb or Cr removal; the result is affected by 
the presence of different solid forms of the metals in the soils, differences in 
pH during extraction, and interference from other cations which complex with 
the chelate (Elliott and Brown, 1989; Brown and Elliott, 1992; Tuin and Tels, 
1990; Hsieh, et ah, 1989; Shirk and Farrel, 1985). The solid forms of Pb or Cr 
present in a soil depend on the source of the contamination and also the extent 
of redistribution of the metal in the soil following contamination. After Pb or Cr 
is added to soil, they may be redistributed by the formation of secondary min- 
eral precipitates, adsorption onto soil mineral particles, or by complexation with 
soil organic matter (Heil, et al., 1996). 

In certain situations, remediation via ex-situ processes may be difficult if a 
site contains utilities or structures. Thus, an in-situ treatment process (i.e., forc- 
ing an extractant through an intact soil to flush out metals) would alleviate some 
of these problems. In general, in-situ technologies are able to treat large volumes 
of soil more economically and more safely than ex-situ technologies because 
there is no excavation (Reed, et al, 1995). However, little fundamental research 
has been carried out on the in-situ flushing of metals (Davis and Singh, 1995). 

Recent soil treatment studies have assessed remediation effectiveness of soils 
spiked with soluble metal salts (Cline and Reed, 1995; Reed, et al, 1995; Davis 
and Singh, 1995; Chen, et al, 1995; Macauley and Hong, 1995). The removal 
efficiencies measured may be greater than those observed when washing cont- 
aminated soils which have been weathered for long periods. In the current study, 
the authors investigated the use of several solutions for Pb and Cr removal from 
a soil contaminated for decades with metals in various insoluble forms. Specif- 
ically, the objectives were to (1) assess the relative abilities of EDTA and NTA 
in the in-situ extraction of Pb and Cr from a contaminated soil and to (2) com- 
pare the effectiveness of in-situ and ex-situ metal removal from the soil. In a pre- 
vious study (Pichtel and Pichtel, 1997), EDTA and NTA solutions were assessed 
for their relative abilities in the ex-situ washing of a Pb- and Cr-contaminated 
soil. Based on these studies, EDTA and NTA concentrations of 0.1, 0.01, and 
0.001 M were selected as soil flushing solutions for the present study. 


The soil material, a mixture of native soil and industrial waste, was collect- 
ed from a closed chemical facility in the United Kingdom. Sample preparation 

Vol. 106 (1997) Indiana Academy of Science 



0.1 M 

0.01 M 


0.001 M 

I 1 I I I 


Pore Volume 

Figure 1. Pb recovery from soil using EDTA at ambient pH (5.5). 


as well as the chemical and physical analyses are described elsewhere (Pichtel 
and Pichtel, 1997). The soil had a pH of 10.3, 1300 mg Pb/kg, and 4940 mg 
Cr/kg. The soil contained 82.8% sand-sized particles, making it suitable for 
soil flushing processes. A sand fraction of 50-80% or more typically increases 
the effectiveness of soil washing (U.S. Environmental Protection Agency, 1991). 

Column studies were conducted using PVC columns measuring 2-cm inter- 
nal diameter and 5 cm in length. Contaminated soil was packed in the column 
with a final bulk density of 1 . 1 g/cm 3 . The column was plugged at both ends with 
rubber stoppers and glass wool. Flushing solutions were introduced from the 
bottom of the columns to saturate the soil. A flow rate of 2.0 ml/min was estab- 
lished for all columns on a Masterllex Model 7568 peristaltic pump. Extraction 
solutions included ethylenediaminetetraacetic acid (EDTA) and nitrilotriacetic 
acid (NTA) at 0.1, 0.01, and 0.001 M. Solutions were used at the ambient pH 
value (5.5 for EDTA and 1 1 . 1 for NTA) or at pH 3.0 using HC1, and 10.0 (EDTA 
only) using NaOH. In one set of experiments, a solution of 0.1 M HO was passed 
through the columns for the first 100 pore volumes with subsequent flushing 
by either 0.1 M EDTA or 0.1 M NTA (ambient pH). 

Column effluent samples were acidified with concentrated HNO. Concen- 
trations of soluble Pb and Cr were measured via flame atomic absorption spec- 
trophotometry (Perkin-Elmer model 2240, Norwalk, Connecticut; Perkin-Elmer. 


Environmental Quality: Pichtel, etal. Vol. 106 (1997) 



T5 70- 

> 60- 

£ 50- 


C£ 40- 


1 H^ 


/ ^^ 


i xt — ^y^ <*— — *^ 

1 /^^ 





1 1 1 1 1 1 1 1 

0.1 M, pH10 
0.1 M, pH 3 

0.01 M, pH 3 

0.01 M, pH10 

100 200 

Pore Volume 

Figure 2. Pb recovery from soil using EDTA at pH 3.0 and 1 1 .0. 

1982). Detection limits of 0.19 and 0.08 mg/L were measured for Pb and Cr, 
respectively, using standards prepared from commercial reagents. 

Comparison of metal levels removed from the soil by EDTA and NTA 
were performed using a one-way analysis of variance and the Student-Newman- 
Keuls Test, if a significant difference was detected (P < 0.05). SigmaStat (ver- 
sion 2.0 on a Windows format) was used for all calculations. 


Lead Removal Efficiencies. Lead removal by 0.1 M EDTA was initially 
rapid (Figure 1). The overall extraction process is consistent with the two-step 
metal desorption process described by Tuin and Tels (1990), Backes, et al. (1995), 
and Cline and Reed (1995); that is, a rapid initial desorption was followed by 
gradual release. The lower extraction efficiency after the first 20-25 pore vol- 
umes is presumably a result of stronger Pb binding to soil solids with decreas- 
ing metal contamination, because binding energies associated with low sorption 
densities are substantial (Benjamin and Leckie, 1981). Additionally, the removal 
of progressively more stable Pb minerals or a slow rate of release of Pb 2+ from 
solid Pb phases may be responsible (Heil, etal, 1996). Reed, etal (1995) mea- 
sured significant Pb removal from contaminated soil columns after only 1-4 pore 
volumes, after which little additional Pb removal occurred. 

After 100 pore volumes, the 0.1 M EDTA solution at ambient pH removed 
71% of the soil Pb, which was significantly greater (P < 0.05) than the 16.8% 

Vol. 106 (1997) Indiana Academy of Science 99 

and 8.0% removed by the 0.01 M and 0.001 M solutions, respectively (Figure 
1). After 200 pore volumes, 89.8% of the soil Pb was extracted with 0.1 M EDTA; 
however, removal at 0.1 M and 0.001 M remained virtually unchanged. Com- 
mon soil metals (e.g., Ca 2+ , Na + , etc.) may compete with Pb for the chelating 
agent so that excess chelant quantities (i.e., well above equimolar concentra- 
tions) are needed to ensure complete contaminant removal (Reed, et al, 1995). 
A chelant concentration of at least 0.00025 M is required for 1:1 concentra- 
tions of Pb:extractant. 

The 0.1 M EDTA solution at pH 3 removed 82.2% Pb after 100 pore vol- 
umes, and 63.3% was recovered with the 0.01 M solution (Figure 2). Heil, et al 
(1996) measured increased Pb removal from three soils as EDTA pH decreased. 
The solubility of many Pb minerals, including Pb(OH)2, PbO, and PbCCh as well 
as other Pb-crystalline solids, will increase as pH is decreased (Lindsay, 1979). 
Protonation weakens the metal-lattice bonds, increasing the dissolution rate. Both 
proton- and ligand-enhanced dissolution mechanisms may be operating simul- 
taneously (Stumm and Wieland, 1990). 

When the EDTApH was increased to 10, the 0.1 M solution removed 84.2% 
Pb after 100 pore volumes, and the 0.01 M solution removed 39.2% (Figure 2). 
At pH values of 1 1 or above, EDTA is present in the completely ionized tetraneg- 
ative form and binds strongly to transition metal cations. EDTA was found effec- 
tive for Pb recovery from soil at high solution pH values in studies by Elliott and 
Brown (1989). Lead solubilization may be partly the result of solubilization of 
soil organic matter or the formation of Pb hydrolysis complexes (Heil, et al, 
1996). Heil, et al. (1996) found, in alkaline solutions, that a high percentage of 
EDTA complexed with Ca and, to a lesser degree, with Mg and other cations. 
The soil in the current study contained 8.2% Ca and 6.0% Mg (Pichtel and Pich- 
tel, 1997); both may compete with target metals for the chelant. The log stabil- 
ity constants for CaEDTA 2 and CaNTA- are 10.7 and 6.4, while the K s for 
MgEDTA 2 and MgNTA" are 8.8 and 5.4, respectively. The K s for PbEDTA 2 and 
PbNTA are 18.0 and 11.3, respectively (Martell and Smith, 1974). 

The present data demonstrates equal or greater Pb extractability in columns 
compared to batch extractions. In batch studies, 0.1 M EDTA was successful, 
both at pH < 4.5 and pH > 12.5, in removing > 90% soil Pb after a single wash- 
ing (Pichtel and Pichtel, 1997). Lead removal efficiencies were high in columns 
compared to batch studies (Reed, et al, 1995). Lead release during column flush- 
ing is apparently enhanced by the higher concentration gradient. 

The quantities of Pb removed from contaminated soils via chelant extrac- 
tion varies. A solution of 0.08 M EDTA removed over 90% of soil Pb from a 
Pb battery-contaminated soil (Elliott and Brown, 1989). Brown and Elliott (1992) 
measured 80% Pb recovery from the same soil at pH 4.0 and 0.04 M EDTA. Tuin 
and Tels (1990) measured variable Pb extraction from contaminated soils using 
EDTA following acidification. 

An increase in NTA concentration resulted in an insignificant increase in Pb 
solubilization (Figure 3). After 200 pore volumes, the 0.1 M NTA solution at 


Environmental Quality: Pichtel, et al. Vol. 106 (1997) 

60 n 


0.1 M, pH11 
0.01 M, pH11 

0.1 M, pH3 
0.01 M, pH 3 


Pore Volume 

Figure 3. Pb recovery from soil using NTA at pH 3 and ambient pH (11.1). 

ambient pH (11.1) removed 50.9% of the soil Pb, which was not significantly 
greater than the 49.5% removed by the 0.01 M solution. The 0.1 M NTA solu- 
tion at pH 3 removed 22.5% after 100 pore volumes and 29.5% after 200 pore 
volumes. A total of 21 .2% was recovered with the 0.01 M solution after 200 pore 
volumes (Figure 3). In batch studies, a range of 12% to 38% Pb was removed 
from this soil in 0.1 M NTA under acidic conditions (Pichtel and Pichtel, 
1997), which was not significantly different from those measured in the cur- 
rent in-situ study. 

Lead recovery in NTA was less than that accomplished by EDTA (Figures 
1-3). Average Pb removed was 87.9% in 0.1 M EDTA (across all pH values) 
compared to 40.2% for 0.1 M NTA. The lower extraction efficiency of NTA 
when compared to EDTA may be due to competition among other soil cations 
(e.g., Ca 2+ ) for the ligands or adsorption of the Pb-NTA complex to the soil sur- 
face. Elliott and Brown (1989) suggest that NTA:Pb ratios greater than 1 : 1 reduced 
Pb recovery because of adsorption of Pb(NTA)1 onto positively charged oxide 
soil components. Additionally, as a result of its weaker complexing ability, NTA 
is less capable than EDTA in preventing Pb hydrolysis under alkaline conditions 
(1989). Castle, et al (1985) found a 10% EDTA solution to be superior to NTA 
in solubilizing Pb; a 90% to 95% removal was measured. In the present study, 
high concentrations of either EDTA or NTA did not reduce metal recovery. 

Vol. 106 (1997) Indiana Academy of Science 


50 n 

0.1 M 


Pore Volume 

Figure 4. Cr removal from soil using EDTA at ambient pH (5.5). 

Flushing the soil columns with HC1 prior to chelant flushing did not signif- 
icantly (P < 0.05) increase Pb-extraction efficiency (data not shown). Tuin and 
Tels (1990) measured 81% and 87% Pb removal from two soils which were first 
extracted by 0.1 M HC1 followed by 0.1 M EDTA. 

Chromium Removal Efficiencies. Overall, Cr removal from the soil was 
substantially lower than that of Pb (Figures 4-5). Tuin and Tels (1990) found 
Cr to be less readily extractable than Pb from four contaminated soils. Assink 
and Rulkens (1987) also measured only slight Cr removal from soil. Metal removal 
is based on the formation of soluble complexes. In the present study, the 
majority of soil Cr is not readily reactive; 89.8% occurs as the insoluble fraction 
(Pichtel and Pichtel, 1997). Therefore, a chelant concentration well above the 
stoichiometric amount is necessary for maximum removal. A minimum chelant 
concentration of 0.004 M is needed to form a 1: 1 ratio of chelant:Cr for opti- 
mal recovery. 

Increasing the EDTA concentration generally resulted in an enhanced recov- 
ery of soil Cr (Figure 4). A total of 40.9% soil Cr was removed with 0. 1 M EDTA 
after 200 pore volumes at ambient pH, and 21.3% and 24.8% was removed at 
0.01 and 0.001 M EDTA, respectively. 

In batch studies, Cr recovery using EDTA at 0.1, 0.01, and 0.001 M was 
maximized at 34-36% up through pH 7 and, in no case, did a single EDTA wash- 


Environmental Quality: Pichtel, etal. Vol. 106 (1997) 

50 n 


Pore Volume 

0.1 M, pH 3 

0.1 M, pH11 
0.01 M, pH 3 

0.01 M, pH11 



Figure 5. Cr removal from soil using NTA at pH 3.0 and ambient pH (11.1). 

ing remove all the soil Cr (Pichtel and Pichtel, 1997). Hsieh, et al. (1989) 
found that Cr removal from a contaminated soil with EDTA ranged from 35- 
83% for 6 to 10 washing cycles, which demonstrates the difficulty of removing 
all adsorbed Cr. Shirk and Farrel (1985) measured only 12% Cr recovery from 
a contaminated soil using EDTA. The poor removal results for Cr may be a result 
of the presence of immobile Cr 3+ species, occurring as oxide compounds (Tuin 
andTels, 1990). 

Chromium removal by 0.1 M NTA (ambient pH 11.1) after 100 pore vol- 
umes was 21.9%, while that by 0.01 M was 1.7% (Figure 5). Beyond pH 10.3 
(i.e., pKa 3 ), the chelant exists as NTA 2 and, to a lesser extent, as NTA 3 , both of 
which allow for partial complexation with the soil Cr. Precipitation as insolu- 
ble oxides is a strongly competing reaction in this pH regime, however. When 
the pH was set at 3.0, Cr removal after 100 pore volumes was essentially unchanged 
with 0. 1 M NTA (20.9%); however, removal increased to 16.9% at 0.01 M. Banat, 
et al. (1974) found negligible Cr solubilization from two river sediments (pH 
6.8 and 7.1) using NTA in batch studies, even after 200 h shaking time. In 
batch tests, Cr removal using 0. 1 M NTA over a range of pH values did not exceed 
14.0% (Pichtel and Pichtel, 1997). 

The stability constant for the formation of CrEDTA" is 17 orders of magni- 
tude higher than that for CrNTA (Martell and Smith, 1974); hence, the EDTA 
should be more effective at removing Cr from the soil. The weaker complexing 
ability of NTA limits its ability to prevent hydrolysis. Furthermore, since all 

Vol. 106 (1997) Indiana Academy of Science 103 

the functional groups of the EDTA are involved in the complexation, little oppor- 
tunity exists for bridging to the soil surface (Elliott and Brown, 1989). Addi- 
tional recovery of Cr at high pH may be limited by the strong affinity of basic 
cations such as Ca and Mg for the chelant. Ca 2+ will strongly bind with the chelant 
at high pH. When the columns were first acidified with HC1, elution by EDTA 
or NTA (ambient pH) did not remove significantly greater quantities of Cr com- 
pared to chelant flushing alone (data not shown). 


Removal of both Pb and Cr from a contaminated, strongly buffered alkaline 
mixture of soil and weathered industrial waste was typically enhanced in col- 
umn studies when EDTA and NTA concentrations were increased. For a non- 
pre-treated (i.e., non-acidified) soil, washing with EDTA above equimolar 
concentrations was more effective than washing with NTA. NTA is less expen- 
sive than EDTA, and less NTA is required, on a weight basis, to form 1 : 1 com- 
plexes with metals. The slow dissolution kinetics of the Pb- and Cr-solid phases 
may deter total metal removal by a chelant, and long-term flushing would be 
needed for complete metal recovery. 

The extent of Pb and Cr solubilization is partly a function of pH. However, 
to completely acidify the soil would prove too costly due to the soil's extensive 
buffering capacity. In addition, high acid strengths destroy soil structure and dis- 
solve much of the soil solids. From a practical standpoint, a soil solution at either 
pH extreme will be corrosive to field washing equipment. The recommended 
washing treatment for the alkaline soil/waste mixture studied, based on pH adjust- 
ment and other practical considerations, was 0. 1 M EDTA without acidification. 

For this study soil, in-situ processes appear to be equally effective as ex-situ 
processes due to the high concentration gradient for soil metals during the flush- 
ing process. Additionally, the particle size distribution of the materials is con- 
ducive to flushing; over 82% of the particles are present in the sand-sized fraction, 
which provides for high permeability and sufficient contact of extracting solu- 
tion with the contaminants. 


Financial support from the Ball State University Energy Center (CERES), 
the Indiana Academy of Science, and the University of Stirling (Scotland) is 
gratefully acknowledged. 


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Elliott, H.A. and G.A. Brown. 1989. Comparative evaluation of NTA and EDTA for extractive decontamina- 
tion of Pb-polluted soils. Water, Air, Soil Pollut. 45: 361-369. 

Heil, D., A. Hanson, and S. Zohrab. 1996. The competitive binding of lead by EDTA in soils and implica- 
tions for heap leaching remediation. Radioactive Waste Manage. Environ. Restoration 20: 111-127. 

Hsieh, H., M. Barnes, and E.Z. Aldridge. 1989. A feasibility study of the removal of chromium from select- 
ed contaminated sites. In: Physiochemical and Biological Detoxification of Hazardous Wastes, pp. 
446-459, Technomic Publ., Lancaster, Pennsylvania. 

Kabata-Pendias, A. and H. Pendias. 1992. Trace elements in soils and plants, 2nd ed. CRC Press, Boca Raton, 
Florida, 365 pp. 

Lindsay, W.L. 1979. Chemical equilibria in soils. John Wiley and Sons, New York, 449 pp. 

Macauley, E. and A. Hong. 1995. Chelation extraction of lead from soil using pyridine-2,6-dicarboxylic 
acid. J. Hazardous Materials 40: 257-270. 

Martell, A.E. and R.M. Smith. 1974. Critical stability constants, vol. 1: Amino acids. Plenum, New York, 
469 pp. 

Perkin-Elmer. 1982. Analytical methods for atomic absorption spectrophotometry. Perkin Elmer, Norwalk, 
Connecticut, 230 pp. 

Pichtel, J. and T Pichtel. 1997. Comparison of solvents for ex-situ removal of chromium and lead from con- 
taminated soil. Envir. Eng. Sci. 14: 97-103. 

Reed, B.E., R.E. Moore, and S.R. Cline. 1995. Soil flushing of a sandy loam contaminated with Pb(II), PbS0 4 , 
PbC0 3 , or Pb-naphthalene: Column results. J. Soil Contamination 4: 243-267. 

Richards, I.G., J. P. Palmer, and PA. Barratt. 1993. Reclamation of former coal mines and steelworks. Else- 
vier, Amsterdam, 718 pp. 

Shirk., J.E. and C.W. Farrel. 1985. Approach to in-situ management of metals. In: Proceedings of the 8' h Madi- 
son Waste Conference, pp. 52-59, Madison, Wisconsin. 

Stumm, W. and E. Wieland. 1990. Dissolution of oxide and silicate minerals: Rates depend on surface speci- 
ation. In: W. Stumm (Ed.), Aquatic Chemical Kinetics, pp. 367-400, John Wiley and Sons, New York, 
545 pp. 

Tuin, B.J.W. and M. Tels. 1990. Removing heavy metals from contaminated clay soils by extraction with 
hydrochloric acid, EDTA, or hypochlorite solutions. Environ. Technol. Let. 11: 1039-1052. 

U.S. Environmental Protection Agency. 1991. Guide for conducting treatability studies under CERCLA: 
Soil washing interim guidance. Washington, D.C., EPA/540/2-9 1/020 A, 38 pp. 

Proceedings of the Indiana Academy of Science 1 05 

(1997) Volume 106 p. 105-111 






John B. Droste and Alan S. Horowitz 

Department of Geological Sciences 

Indiana University 

1005 East 10th Street 

Bloomington, Indiana 47405 

ABSTRACT. Electric logs and samples of well cuttings indicate the existence 
of limestone bodies in the lower and middle Brazil Formation of southwest- 
ern Indiana and western Kentucky. The limestone bodies are elongated north- 
east-southwest, are as much as 60 feet thick, are as much as 3 miles in length, 
are interpreted to have formed on slight elevations on the sea floor, and were 
accumulated contemporaneously with adjacent terrigenous muds and sands 
until they were finally smothered by these terrigenous sediments. 

KEYWORDS: Brazil Formation, limestone, Pennsylvanian, subsurface south- 
western Indiana, subsurface western Kentucky. 


The Brazil Formation in Indiana lies below the Staunton Formation and 
above the Mansfield Formation in the Raccoon Creek Group (Figure 1). In the 
type area (Brazil, Clay County, Indiana), the Brazil includes rocks from the base 
of the Lower Block Coal Member to the top of the Minshall Coal Member (Hutchi- 
son, 1976). The Upper Block Coal Member is the only other named member of 
the Brazil Formation. The Brazil coals have irregular distributions along the out- 
crop and very limited distributions in the subsurface. Because the Minshall coal 
is of limited extent, the stratigraphic position of the base of the more widespread 
Perth Limestone Member, or of the sandstone that replaces it, was used in an 
earlier subsurface study (Droste and Horowitz, 1995) to mark the top of the 
Brazil. This "working" definition of the top of the Brazil is used in the present 
study. Following conventional usage, the base of the Brazil is placed at the strati- 
graphic position of the base of the Lower Block Coal. 

This study expands geographically the earlier report of limestones in the 
Brazil Formation in Vanderburgh County, Indiana (Droste and Horowitz. 
1995) by including all the subsurface area of southwestern Indiana and western 


Geology & Geography: Droste and Horowitz Vol. 106 (1997) 






PERMIT #21529 

Figure 1 . An electric log from a well in 
Vanderburgh County, Indiana, showing 
thick limestone in the Brazil Formation 
and the stratigraphic nomenclature used 
in this report. The position of the well 
is shown in Figure 4 at location 4C. 


Perhaps the only significant mention 
of limestone in Brazil rocks at the surface 
is the report of the fusulinid genus Fusu- 
linella in an unnamed limestone between 
the Lower and Upper Block Coals in Clay 
County, Indiana (Shaver, 1981). The rather 
common occurrence of limestone in the 
Brazil in the subsurface initially appeared 
to be limited to Indiana. However, the 
authors have found "Curlew" (referring to 
the Curlew Limestone Member of the Trade- 
water Formation) on electric logs in Ken- 
tucky that are assigned to any of several 
limestone intervals in rocks equivalent to 
the upper part of the Brazil Formation of 
Indiana. One to three intervals of lime- 
stone, each less than 10 feet thick, are pre- 
sent in numerous wells in southwestern 
Indiana and western Kentucky. These thin 
intervals of limestone in the upper Brazil 
are principally wackestones that in places 
contain chert. Packstones are less abun- 
dant and typically are not cherty. 

This report focuses on thick limestone 
intervals in the middle and lower part of 
the Brazil. Samples (well cuttings at 10- 
foot spacing) in the interval of high resis- 
tivity in the Brazil Formation in the Kamp 
and Rollett #2 well (Figure 1) record almost 
60 feet of limestone, which is the thickest 
limestone interval for which samples were 
available for this study. The sample stud- 
ies have been restricted to materials from 
the Brazil interval in wells on file at the 
Indiana Geological Survey. Samples from 
wells in Kentucky were not examined. 
Samples, typically taken at 10-foot inter- 
vals, from many wells in southernmost 
Vanderburgh County (Droste and Horowitz, 
1995) form the basis for the following 
description of thick limestones in the Brazil 

Vol. 106 (1997) Indiana Academy of Science 







A s "* 


















c s <^ 



i i 


1 T 

1 C 







Figure 2. Typical electric logs from wells in Indiana and Kentucky that show limestone 
intervals from 20 to more than 40 feet thick. The positions of the wells are shown in Fig- 
ure 4 at locations 2A, 2B, 2C, and 2D. 

The biotic skeletal components of the limestone are primarily echinodermal 
fragments, bryozoans, and brachiopods, a typical late Paleozoic faunal associa- 
tion interpreted as living on shallow marine shelves. Other minor faunal com- 
ponents identified in sample fragments are mollusks, ostracodes, and trilobites. 
The sample from the basal 10 feet of the limestone contains mainly wacke- 
stone with lesser amounts of packstone. This interval is locally cherty, suggest- 
ing the presence of silica-bearing organisms, perhaps sponges. The next higher 
10-foot interval shows a much reduced abundance of wackestone and a greatly 
increased abundance of packstone. Chert is seldom encountered in samples of 


Geology & Geography: Droste and Horowitz Vol. 106 (1997) 





-J i l - 

r- 1 c 

CL * Y I I > 







/ £— G,B !2i^; DUBOIS |_ _ 

\ ^ fXPsf^ •:•'• /*^ ,v, [crawforI 

wapitc. / S'-v ■•.'•'• //■ . \ 

J- -I 

-^J WARRICK L .^ 1 


'0 M;& 

k 1 ' HANCOCK I ' a 


A /breckin-" 






KINS l yS? 

^ ^ ^ , MUHLENBERG « P / ~f~ 

) ^LOWELL i ^^k.\% \ A 

.mccracken^U / L ™ v / ^.^ \ ^_^>_ Wen" 

6 l 7 I 8 \ 9 I 10 I 11 I 1 2 Th I 14 MsYlfc i 17 I 18 I 19X201 21 Al I 23 1 24 1 25 1 26 I zj 1 28 1 29 I to 1 31 I 32 I 33YW3S I 36 I tl 


1 .' I I 

30 Miles 

50 Km 

Figure 3. A map showing the distribution of thick limestone intervals in the Brazil For- 
mation. Stippling indicates areas within which at least one well per section (square mile) 
contains an interval of limestone 10 or more feet thick. Heavy dots show locations of 
sections in which at least one well has a limestone interval 20 or more feet thick. 

the second 10-foot interval. In the sample of the third 10-foot interval, grain- 
stone is almost as abundant as packstone. Limestones more than 30 feet thick 
are composed mainly of grainstone and have the appearance of typical late Mis- 
sissippian limestone. However, oolites, common in numerous late Mississippi- 
an limestones, have not been observed in the limestones in the Brazil Formation. 


The unusually thick limestone in the Brazil interval in southernmost Van- 
derburgh County (Figure 1) is an uncommon occurrence. Several electric logs 

Vol. 106 (1997) Indiana Academy of Science 




sec. 3 

sec. 2 





yy* 7 / 


(J/ ■ 


sec. 10* 

• sec.1 1 


I .sec.6\ 

Cl = 10' 



sec. 17 

Cl = 20' 

sec. 16 

sec. 21 


!L _ [i M t 


Figure 4. Maps showing the northeast-southwest alignment of the thick limestone bod- 
ies in the Brazil Formation of Vanderburgh County (A, C) and Posey County (B), Indi- 
ana. Contour interval as specified; dots indicate locations of well log control. For the 
location of these areas, see the index map. 

from wells in other locations show the more typical thickness of Brazil lime- 
stone bodies (Figure 2). The thick limestone intervals in the two wells from Ken- 
tucky (Figure 2C and 2D) and from southernmost Vanderburgh County, Indiana 
(Figure 1), are in the lower Brazil. At two locations farther north in Indiana (Fig- 
ures 2A, 2B), the thick limestone interval is in the middle Brazil. The environ- 
ment suitable for limestone deposition clearly is younger in the more northerly 
locations in Indiana. 

Limestone intervals ten or more feet thick in the Brazil Formation have a 
greater areal distribution in southern Indiana than in western Kentucky (Figure 
3). The stippling in Figure 3 indicates areas where at least one well per section 
(square mile) contains an interval of limestone 10 feet or more thick, but with- 
in the shaded areas, the thicker limestone intervals are discontinuous even with- 
in a single section. The heavy dots (Figure 3) within the stippled areas mark 

1 10 Geology & Geography: Droste and Horowitz Vol. 106 (1997) 

locations of sections where at least one well contains an interval of limestone 20 
or more feet thick. 

In several areas in Indiana, closely spaced wells with geophysical logs per- 
mit mapping of the distribution of single bodies of thick limestone. Two loca- 
tions in Vanderburgh County and one location in Posey County (Figure 4) 
show the shape and orientation of the limestone accumulations. The limestone 
deposits are elongate in a northeast-southwest direction. These maps clearly illus- 
trate, as noted above, that the thick limestone intervals are discontinuous with- 
in the same section. 

The limestone intervals in the upper Brazil are thinner and more continuous 
than the thick limestone intervals in the middle and lower Brazil. 


We can only speculate about the environmental conditions that permitted the 
formation of the thicker limestone accumulations. Shallow marine shelf depo- 
sition was common to this area during the Mississippian Period and, at times, 
apparently continued into the Pennsylvanian based on the limited biotic evidence 
available. The fauna also indicates marine waters, and the abundance of echin- 
odermal debris, presumed to be principally crinoidal plates, suggests that crinoidal 
thickets existed. These thickets would have needed a suitable substrate on which 
to grow and, in calm weather, sufficient energy in the form of waves and cur- 
rents to bring in nutrients but not to carry sufficient terrigenous sand and mud 
to smother the thickets. Kvale, Furer, and Mastalerz (1996) have reported tidalites 
above the Lower Block Coal Member at the outcrop and in the shallow subsur- 
face in Daviess County, Indiana. This confirms the presence of active tidal cur- 
rents during Brazil time no more than 30 kilometers from the nearest subsurface 
carbonate buildup discussed here. The thickets are presumed to have been above 
storm wave base and probably would have been swept free of some accumulat- 
ed carbonate or terrigenous debris during storms. The biotic debris is believed 
to have been produced locally as evidence is lacking for distant transport of skele- 
tal debris to the site of deposition. Electric log correlations suggest contempo- 
raneous terrigenous sands and muds accumulated adjacent to the loci of limestone 

The suggested environment of deposition for the thick limestone accumu- 
lations is as follows. Slight elevations on a shallow marine shelf permitted the 
establishment and growth of crinoid thickets above storm base. The accumula- 
tion of biotic debris accentuated the relief on the sea floor and permitted skele- 
tal organisms to flourish. Accumulation continued until a change in currents or 
other water conditions led to smothering of the limestone bottoms by terrige- 
nous debris. The northeast-southwest orientation of the elongate deposits may 
be the result of very minor tectonic control or slight variations in the rates of 
subsidence from area to area on the sea floor. 

Vol. 106 (1997) Indiana Academy of Science 111 


The existence of thick normal marine limestone accumulations in the Brazil 
Formation clearly documents a strong marine influence in the environments of 
deposition in southwestern Indiana and western Kentucky during Brazil time. 
Crinoid thickets apparently colonized slightly elevated sites on the sea floor that 
were swept by waves and currents sufficient to bring in nutrients but lacking 
smothering terrigenous materials. Basal wackestones grade upward into pack- 
stones. In deposits 30 or more feet thick, the upper grainstones have the 
appearance of the better-known shallow water limestones of the underlying Mis- 
sissippian rocks. The Brazil limestones were laterally contemporaneous with and 
were ultimately covered by terrigenous debris. 


The authors thank J.R. Dodd, H.H. Gray, S.J. Keller, and N.G. Lane for the 
critical review of several manuscript versions of this report. 


Droste, J.B. and A.S. Horowitz. 1995. Unusual carbonate buildup in the Upper Mansfield and Brazil Forma- 
tions, Vanderburgh County, Indiana. Indiana Acad. Sci. 111th Annu. Meeting, Progr. Abstr., p. 70. 

Hutchison, H.C. 1976. Geology of the Catlin-Mansfield area, Parke and Putnam Counties, Indiana. Indiana 
Geol. Surv. Bull. 54, 57 pp. 

Kvale, E.P, L.C. Furer, and M. Mastalerz. 1996. Exploration models for selected economic coals in south- 
eastern Daviess and southwestern Martin Counties. Indiana Geol. Surv. Open-File Rep. 96-2, 28 pp. 

Shaver, R.H. 1984. Atokan Series concepts with special reference to the Illinois Basin and Iowa. Oklahoma 
Geol. Surv. Bull. 136: 101-113. 

Proceedings of the Indiana Academy of Science 113 

(1997) Volume 106 p. 113-134 

HERMAN T. BRISCOE (1893-1960): 




Harry G. Day 

Department of Chemistry 

Indiana University 

Bloomington, Indiana 47405 

ABSTRACT: Herman Briscoe was so outstanding in the Shoals, Indiana, pub- 
lic schools that after he finished high school in 1912 and took teacher train- 
ing courses that summer at Indiana University, he taught in Shoals High School 
for the next six semesters. At the same time, he took summer courses at Indi- 
ana University. He completed an A.B. degree in chemistry with high distinc- 
tion in 1917. Over the next five years, he served consecutively and briefly as 
Superintendent of Schools in Shoals, then enrolled in the Officers Training 
Program in the U.S. Army, was transferred to industrial work on chemical 
explosives, and, after the war, devoted brief periods of time to teaching chem- 
istry at Stark's Military Academy, Harvard University, and Colby College 
before returning to Indiana University for graduate work in 1922. He joined 
the faculty at Indiana University in 1924, immediately after receiving his Ph.D., 
and became Professor of Chemistry in 1928. From 1925 to 1942, he directed 
the graduate work of 25 students, seventeen of whom earned a Ph.D. He was 
also the author or coauthor of 23 scientific papers and one U.S. patent. Between 
1931 and 1951, he published several very successful chemistry textbooks. His 
standing as a chemistry teacher and public lecturer rose correspondingly. With 
the appointment of Herman B Wells as Acting President of Indiana Univer- 
sity in 1937 and President in 1938, Briscoe was promptly selected, along with 
two other well-regarded faculty members, to serve on Wells' Self-Survey Com- 
mittee. Many sound and basic changes in the University were recommended 
by the Committee, and within a few years, all the proposed changes were 
adopted. Early in this notable transformation, Briscoe became the first Dean 
of the Faculties. Soon, new responsibilities included his appointment as the 
first Vice President of the University. Functioning closely together as an admin- 
istrative team, Briscoe and Wells became recognized as great role models. 
When Briscoe retired (1959), the most meaningful tribute to his excellence 
was made by Wells (Gaugh, 1959): Briscoe was the "wisest educational admin- 
istrator in America." Following Briscoe's untimely death in 1960, a major res- 
idence complex was named after him, and the endowed Briscoe Professorship 
in Chemistry was established. Both serve as tributes to Dr. Briscoe as a role 
model in chemical education and academic administration. 

KEYWORDS: Chemical education, chemistry as a changing science. Her- 
man T Briscoe, research is second in value only to teaching, structure and 
properties of matter, student guidance is basic in education, Herman Wells. 

Residents of Indiana, especially members of the Indiana Academy of Sci- 
ence, can gain much satisfaction from learning about Herman Thompson Briscoe 

1 14 History of Science: Day Vol. 106 (1997) 

(1893-1960). His influence in chemical education and various aspects of uni- 
versity administration as well as his character and ability to guide others in diverse 
matters was far greater than is commonly realized. The dimensions of Dr. Briscoe's 
professional life are extensively cited in The Development of Chemistry at 
Indiana University 1829-1991 (Day, 1992) and formed the basis, along with 
other personal and professional appraisals of his life and contributions, of an 
unpublished biographical sketch prepared by the author in the 1980s (Day, 1987). 


Dr. Briscoe was born and raised on a farm near Shoals, Indiana. From his 
earliest years, he evidenced exceptionally high intelligence and pleasing per- 
sonal qualities. In the modest Shoals High School, he consistently earned high 
grades while completing the following units of credit: English composition 
and literature 3, Mathematics 3, Latin 4, German 3, History 1, and Geology x h. 
After graduation, he took teacher preparation courses in the 1912 summer ses- 
sion at nearby Indiana University and received a Class A certificate for teach- 
ing. Beginning immediately, and during the following three academic years, he 
taught several courses — including Latin — at Shoals High School. Before the end 
of the 1914-15 school year, he had become the school's Principal. During the 
summers of 1913 and 1914, he took additional courses at Indiana University. 

Briscoe returned to Indiana University on a full-time basis in the summer of 
1915, and in October 1917, he received an A.B. degree in chemistry with High 
Distinction. In addition, he was elected to membership in Phi Beta Kappa. 
During his two years as a full-time university student, he was active in the strong 
debating program, which competed with Butler, DePauw, and Earlham. Wendell 
L. Willkie (1892-1944) was a fellow member of the debate group. In his sec- 
ond full year, Briscoe was President of the University's Indiana Debating League. 
Among other responsibilities, he was a member of the Indiana Club which empha- 
sized the "ideals of high scholarship, and ambition for proficiency in dramatic 
attainments" (Day, 1987, p. 2). 

At the end of the 1917 summer term, the 24-year-old chemistry graduate 
became Superintendent of Schools in Shoals! Briscoe served in this role until he 
enlisted as a private in an Officers Training Program in the U.S. Army in May 
1918. The Army transferred him to the Hercules Powder Company in Ohio, where 
he remained, working on chemical explosives, until February 1919. From then 
until 1922, the readjusting former soldier held successive teaching assignments 
at Stark's Military Academy in Alabama, Harvard University, and Colby Col- 
lege in Maine. Finally, he returned to Indiana University to do his graduate work 
in chemistry (1922-1924). Under the guidance of Professor Frank C. Mathers, 
he received A.M. and Ph.D. degrees in chemistry. 

His 149-page substantive and exceptionally well- written doctoral thesis was 
entitled "The Properties of Dolomitic Limes as Related to the Properties of the 
Stones, the Conditions of Burning, and Subsequent Treatments." The principal 
findings of his research were published in the 1925 issue of the Proceedings of 

Vol. 106 (1997) Indiana Academy of Science 1 15 

the Indiana Academy of Science. In 1926, a U.S. patent (1,588,253) was issued 
to Briscoe and Mathers covering some of his thesis work — a process for pro- 
ducing plastic dolomitic limes (Day, 1987, p. 3). In 1927, more of their research 
on the plasticity of finishing limes was reported in a journal published by the 
American Chemical Society, Industrial and Engineering Chemistry. This early 
focus on research in the laboratory was influential in shaping the substance and 
diversity of Briscoe's scientific interests and understanding. 

Mounting teaching responsibilities — starting even before the completion of 
his doctoral work in 1924 — diminished the time available for Briscoe's research 
activities. However, from 1925 to 1942, Briscoe supervised the graduate-level 
research of 25 different students, and 17 received Ph.D. degrees in chemistry 
from Indiana University. Most of the research was on the conductivity, chemi- 
cal reactions, and physical properties of substances in nonaqueous solvents. 
Briscoe also authored or coauthored 23 publications: seven were in the Pro- 
ceedings of the Indiana Academy of Science, nine in the Journal of Physical 
Chemistry, and the remainder were in various other journals. Two papers were 
concerned with teaching new concepts on acids and bases, and two others were 
on manpower management problems in World War II. 


1925. The calcination of dolomitic limestone. Proc. Indiana Acad. Sci. 35: 

1927. (With EC. Mathers). Plasticity of finishing limes. Ind. Eng. Chem. 19: 

1929. (With H. Hunt). The conductivity of solutions of some aliphatic acids in 

water and ethyl alcohol. J. Phys. Chem. 33: 190-99. 
1929. (With H. Hunt). The electrical conductivity of organic acids in water, alco- 
hols and acetone and the electronic structure of the acids. J. Phys. Chem. 

33: 1495-1513. 
1929. (With H. Hunt). Factors determining electrolytic dissociation. J. Chem. 

Ed. 6: 1716-1725. 
1929. (With F.M. Whitacre). Esterification in presence of anhydrous salts. Proc. 

Indiana Acad. Sci. 38: 187-194. 
1932. (With F.J. Welcher). The analysis of anions. Chem. News 145: 161-170. 

1932. (With P.E. Williams). The separation and identification of alkaline earths. 
Chem. News 145: 177-184. 

1933. (With W.L. Bright). The acidity of organic acids in methanol and ethyl 
alcohol. J. Phys. Chem. 37: 787-796. 

1933. (With F.E. Dolian). Vapor phase esterification. Proc. Indiana Acad. Sci. 
42: 101-107. 

1935. (With F.E. Dolian). The refractive indices of non-aqueous solutions of 
metallic chlorides. Proc. Indiana Acad. Sci. 45: 110-115. 

1936. (With R.C. Gore). The dielectric constants of solutions of some organic 
acids in ethyl alcohol and benzene. J. Phys. Chem. 40: 619-625. 

1 16 History of Science: Day Vol. 106 (1997) 

1937. (With KM. Whitacre). Esterification in the presence of anhydrous salts. 
Proc. Indiana Acad. Sci. 46: 133-141. 

1937. (With F.E. Dolian). The viscosities of solutions of chlorides in certain sol- 
vents. J. Phys. Chem. 41: 1129-1138. 

1938. (With T.P. Dirkse). The electrodeposition of metals from non-aqueous solu- 
tions. Metal Ind. (New York) 36: 284-285. 

1938. (With J.S. Peake). Measurement of ionization constant of benzoic acid 
using silver chloride electrodes. J. Phys. Chem. 42: 637-640. 

1939. (With W.H. Cathcart and R.H. Treadway). The bromination of acetone in 
non-aqueous solutions. Proc. Indiana Acad. Sci. 48: 92-97. 

1940. (With T.P. Dirkse). The conductance of salts in monoethanolamine. J. Phys. 
Chem. 44: 388-397. 

1940. Teaching the new concepts of acids and bases in general chemistry. 

J. Chem. Ed. 17: 128-130. 
1942. (With J.T. Pinkston). Conductometric titrations in non-aqueous solutions. 

J. Phys. Chem. 46: 469-473. 

1942. (With W.T. Rhinehart). Relative viscosity of non-aqueous solutions. 
J. Phys. Chem. 46: 387-394. 

1943. Training program for chemists. Chem. Eng. News 21: 1702-1704. 

1944. The crisis in chemical manpower. Chem. Eng. News 22: 584-587. 


The author received many helpful letters from people who knew Briscoe 
very well during his student years and/or other periods of his later life. The author 
also had conversations with most of those who wrote to him. 

Russel L. Hardy (A.B. '22 and A.M. '23) wrote in 1962 (Day, 1987, p. 40) 
that ". . . he [Briscoe] was born with intellectual honesty and this hereditary fac- 
tor governed his life." Also, in this long and most sincere letter, he concluded 
that, "Herman's . . . passion for an honest, thorough intellectual performance 
by his teachers ... is no accident nor surprise that it emerged as the dominating 
factor in his thinking and able planning for Indiana University during the remain- 
ing years of his life." At the time of his retirement in 1962, Hardy was Director 
of Electrochemical Production for the DuPont Company 

Hardy, Mrs. Briscoe (formerly Orah Cole), and Mrs. W.J. Sparks (Mered- 
ith Pleasant) provided considerable information about the swift and unexpect- 
ed marriage of Herman and Orah. As Hardy (on file in the Chemistry Archives) 
commented, "He [Briscoe] seemed to be almost totally preoccupied by work" 
and he ". . . had no personal interest in any young lady at the time." But in May 
1928, Meredith Pleasant, a chemistry graduate student, arranged for Orah, her 
roommate, to have a blind date with the young Associate Professor. Later, Mered- 
ith married a fellow graduate student, William J. Sparks, who eventually became 
one of the most highly recognized chemists in America. But the romantic fire 
she ignited through the blind date swept on and three months later Herman and 
Orah were married. During the brief courtship, Herman was elevated to the rank 

Vol. 106 (1997) Indiana Academy of Science 1 17 

of Professor of Chemistry. In a letter to the Sparks written in 1960 shortly after 
her husband's death, Orah stated: "Meredith, that blind date you arranged for us 
was back in May 1928. We never forgot who brought us together." The Briscoes 
had four children: Mrs. Stephen G. Ayers (Catharin Alice), Robert Herman, 
William Cole, M.D., and James Frederick (deceased in early childhood). Each 
surviving child married and had children. 


Briscoe's great interest in and effectiveness at teaching chemistry were reflect- 
ed in the chemistry texts and laboratory manuals that he authored between 1927 
and the early 1950s. His initial effort was a set of mimeographed notes on 
qualitative chemical analysis which focused on "the theories and principles of 
electrolytic solutions and the properties of the cations and anions which are 
involved in the various analytical procedures" (Briscoe, 1931, p. iii). 

In his textbooks, enough substantive historical background was lucidly woven 
into the presentations to provide the student with both perspective and under- 
standing. For example, Briscoe wrote "the path of thought which was followed 
during the centuries of alchemy and the dark ages did not lead toward truth 
. . ." (Briscoe, 1931, p. 38). To help the students understand more about nature 
through modern chemistry, he wrote (Briscoe, 1938, p. 38): 

Perhaps they [the alchemists] erred no more than we err at present in follow- 
ing rather blindly a path of thought and reasoning which would lead to a cross- 
roads of uncertainty if we should follow it back to its origin. From this crossroads 
different routes lead out into darkness of the unknown, and, perhaps, only one 
of them leads to truth, which is always the ultimate goal. 

In this vein, Briscoe elaborated on searching for paths leading to truth through 
the methods of science. Briscoe (1938, p. 38) referred to: 

John Dalton and the scientific world of his day [who] stood at such a cross- 
roads. They were confronted with the question of the structure of matter. Dal- 
ton proved that matter consists of "atoms" by certain experiments in analysis 
which established the Law of Multiple Proportions . . . Soon after the theory 
was announced, knowledge of atomic weights began to accumulate. . . . 

The first 142 pages of Briscoe's 279-page book were devoted to theories and 
general principles; the remainder of the book provided informative instruction 
on procedures along with clarifying explanations and stimulating questions. The 
book emphasized Briscoe's belief (1938, p. 38): 

. . . that laboratory work proves its worth forces him, to investigate for him- 
self and to explain the results of his investigations in terms of his own knowl- 
edge and experiences. 

Also of note was Briscoe's widely read 420-page book. The Structure and 
Properties of Matter, which was published in 1935. Briscoe started teaching this 

118 History of Science: Day Vol. 106 (1997) 

subject in 1925. In 1934, while the book was in press, Robert E. Lyons, Profes- 
sor and Head of the Department of Chemistry, described the forthcoming book 
as "a splendid interpretation of what is now known concerning the structure of 
matter in terms of the chemical behavior of matter" (Day, 1987, p. 4). This book 
was the first publication from the Chemistry Department that compared favor- 
ably with other first-class scientific books. 

In the same year (1935), Briscoe's General Chemistry for Colleges appeared. 
In 1937, he published a related treatment on general chemistry, An Introduc- 
tion to College Chemistry, the fourth edition of which appeared in 1949. Fol- 
lowing Dean Briscoe's death in 1960, William Briscoe discovered that his father 
had almost completed either another revision of one of his textbooks or an entire- 
ly new chemistry textbook. 

Briscoe's philosophy of teaching was reflected within the preface to the last 
edition (4th) of General Chemistry for Colleges (Briscoe, 1949, p. viii): 

Instead of displacing the subject matter of what we may call classical chem- 
istry, new information and new theories are discussed alongside the old. . . . 
If this practice serves no other purpose, it will help the student see that 
chemistry is a changing, developing science and, perhaps, will cause him to 
realize the possibilities that lie ahead of us [as] we extend and revise our 
present knowledge of the subject. 

Even before the highly fruitful H.B Wells - H.T. Briscoe combination in aca- 
demic planning and administration became acknowledged, Briscoe had gained 
recognition for his excellence in teaching chemistry as well as for the high qual- 
ity of his chemistry textbooks (Wells, 1980). Many tributes emphasizing his inter- 
est in teaching were written following Briscoe's untimely death. Especially 
meaningful was the letter written by Frank J. Welcher (A.B. '29, Ph.D. '32), 
whose doctoral research was supervised by Briscoe: "He had the teacher's voice, 
the teacher's care of preparation, and above all, the teacher's passionate inter- 
est that students should fulfill themselves" (Day, 1987, p. 7). Perhaps the epito- 
me of summations is in the measured words of F.T. Gucker, then Dean of the 
College of Arts and Sciences at Indiana University (Day, 1987, p. 7): 

When I came to Indiana University 14 years ago [1947] as chairman of the 
Department of Chemistry I knew Herman T. Briscoe through his books on 
general chemistry and on the structure and properties of matter. Like many 
others throughout the country who used these books in their classes, I admired 
the clear, lucid, and interesting style in which he presented chemical facts and 

The highly respected Gucker served as Chairman of the Department of Chem- 
istry for four years before he became a Dean, a position he held until his age 
necessitated retirement from administrative responsibilities. One section of 
Briscoe Quadrangle (a dormitory complex) is now named in memory of Dean 
Gucker, who died in 1973. 

Vol. 106 (1997) Indiana Academy of Science 119 


Dr. Briscoe accepted scores of invitations to address various local and other 
largely in-state groups and organizations. The topics were substantive, and 
they were usually concerned with chemistry or science in general. A number of 
examples should suffice to show the breadth and scope of these talks. In 1925, 
he spoke to the Bloomington High School Science Club on "The Structure of 
the Atom." In 1928, he spoke before the Physics and Chemistry Sections at the 
Indiana State Teachers' Convention in Indianapolis on "The Electron Theory 
of Valence." "Atomic Theories" was the topic of his talk to the Indiana Univer- 
sity Physics Club in 1933. In 1934, he addressed the campus YWCA on "Con- 
flicts between Science and Religion." According to the campus newspaper (Indiana 
Daily Student), Briscoe "thinkfs] there exists no conflict between science and 
religion." In 1934, he participated in a symposium on "Teaching Methods in 
General Chemistry" at a regional meeting of the American Chemical Society 
in Louisville. In 1936, he spoke at the annual meeting of the Indiana State 
High School Chemistry Teachers' Association at Purdue. His topic was "Caus- 
es of Students' Difficulties in Chemistry." In 1937, under the auspices of the 
Indiana Section of the American Chemical Society, his topic was "Cosmic Rays" 
before a science club meeting at Hanover College. In 1938, as the new Chair- 
man of the Department of Chemistry, he presented a paper at a Bloomington 
meeting of the Indiana Section of the American Chemical Society on a new per- 
spective on "The Place of Chemistry in Indiana University." In 1939, he spoke 
twice at the Indianapolis Center of the Extension Division. The topics were "New 
Products of the Chemical Industry" and "Chemistry under Nazism." 


Soon after Herman B Wells became Acting President of Indiana University 
on 1 July 1937, he instituted a planning process by establishing a Self-Survey 
Committee. On January 15, 1938, the Trustees approved the appointment of three 
highly regarded faculty members to the committee — Herman T. Briscoe, Pro- 
fessor of Chemistry and a dedicated member of the College of Arts and Sciences; 
Wendell W. Wright, Professor of Education; and Fowler V Harper, Professor 
of Law. Harper was designated Chairman, and Briscoe was Secretary. Of the 
three, Briscoe was the only alumnus of Indiana University. Wells stated in his 
autobiography (1980, p. 97) that "Wright was conservative, Briscoe was mod- 
erate, and Harper was liberal." All three worked together efficiently and with 
great thoroughness, understanding well the high importance of their assignment. 
No diminution in teaching load and related academic duties was noted for Briscoe 
and presumably the same applied to the other two Committee members. 

Naturally, the Committee solicited the views of faculty members and oth- 
ers, both locally and nationally, on the best objectives for the University. As Clark 
(1973, p. 372) said; 

120 History of Science: Day Vol. 106 (1997) 

The organizational chart projected by the survey committee was a study indeed 
in a step from rigid simplicity and personal control to one of complexity. In 
fact the committee's charted proposal was more a prescription for revolu- 
tionizing the administrative system of the university than anything of the 
sort had been since the founding of the institution. Not only was this branch 
of university government to be updated, but it was to be brought into proper 
relationship with both the institutional and professional aims and objectives 
set forth in the general program. 

In addition to proposing major administrative changes, considerable atten- 
tion was given to faculty development and to strengthening both teaching and 
research. The Committee emphasized that Indiana University was far behind 
other Big Ten Schools in its level of support for faculty research by pointing out 
that in 1936-37 Indiana University spent only 1.7 percent of its total budget in 
support of original research as compared to an average of more than 10 percent 
at 135 publicly-controlled universities listed in the biennial survey of the U.S. 
Department of Education. The Committee also stated in their report that a marked 
increase in the effectiveness of the University would require a rigorous and com- 
prehensive program of faculty recruitment to attract and hold the most promis- 
ing talent in teaching and research. 

One of the major recommendations of the Committee was to establish the 
office of Dean of the Faculties. The recommendation was promptly approved. 
The first Dean was Dr. Briscoe. On May 31, 1940, President Wells sent a mimeo- 
graphed letter to every faculty member announcing the appointment effective 
the next day. The specific responsibilities of the Dean of the Faculties were 
listed as follows (Day, 1987, p. 21): 

1 . He will share with me general responsibility for the academic adminis- 
tration of the university and will receive and act upon all academic pro- 
posals and problems during my absence. 

2. He will share with me responsibility for public appearances. 

3. He will be an ex officio member of the general university standing and 
special committees unless at the time of the appointment of the com- 
mittee, it is specifically stated that he will not serve. 

4. He will be a member of all Faculties of the University, as is the Presi- 

5. He will assume from time to time such additional responsibilities as may 
seem desirable. 

6. He will continue to be in charge of the guidance program [This evolved 
from the Self-Survey Committee in May 1939. As stated by Clark (1977, 
p. 33): "Herman Briscoe almost single-handedly devised an advisory sys- 
tem which supplanted an older plan and prepared the way for institut- 
ing the Junior Division [now University Division]."] 

7. He will continue as Chairman of the Department of Chemistry until a 
successor can be selected, seeking such relief there at present as possi- 
ble for him to effect. 

Many laudatory expressions of approval to Briscoe's appointment were imme- 
diately forthcoming. For example, Lee Norvell, then Associate Professor of Eng- 
lish and Radio Director, wrote: "I honestly believe that his appointment will 

Vol. 106 (1997) Indiana Academy of Science 121 

receive the unanimous approval of the faculty. I know of no one else of whom 
this could truthfully be said" (Day, 1987, p. 21). In 1941, Briscoe's title was 
changed to Vice President, Dean of the Faculties. This change met with equal- 
ly strong approval. 

The Self-Survey Committee began deliberations early in 1938. By Decem- 
ber 1939, Briscoe, Wright, and Harper had presented their full report to the fac- 
ulty for thorough study and debate. In his comprehensive analysis of the report 
and its effects on the University, Clark (1973, p. 382) wisely concluded: 

For the university the immediate result of the self-study was the aid it 
rendered in a realignment of the institution to recognize new educational oppor- 
tunities and to meet the needs of the near future. The luck of time and histo- 
ry favored Indiana University in 1940. Its administration and a major 
portion of its faculty no doubt became aware of the currents in post-Depres- 
sion America of educational reform and were emotionally and intellectually 
prepared to accept change with little loss of momentum or desecration of 

If Dr. Briscoe had not been afflicted with serious and lingering health 
problems (he had a cerebral hemorrhage in 1945, when he was only 52), many 
individuals have speculated that his remaining contributions would have been 
substantially greater, but his role on the Self-Survey Committee and in the adop- 
tion and implementation of its recommendations were scarcely surpas sable. 


On March 22, 1938, Acting President Wells became President of the Uni- 
versity. During his first nine months of service, major planning and systematic 
changes toward restructuring and energizing the institution, particularly in 
matters of academic organization and higher goals, were implemented. In August 
1938, Dr. Lyons was almost 69 year of age, and he had been Head of the 
Chemistry Department since 1895. "On 4 August 1938 Dr. Robert E. Lyons sub- 
mitted his letter of resignation as Professor of Chemistry and Head of the Depart- 
ment of Chemistry effective August 11, 1938 " (Day, 1992, p. 250). In his 

brief and courteous letter, Lyons strongly recommended that his successor should 
be Dr. Briscoe. The resignation and the recommendation were accepted by the 
Trustees on August 13. The making of an essentially new department began when 
Briscoe was designated Chairman. 

Besides this new responsibility, his regular teaching duties, and his pressing 
service on the Self-Survey Committee, more "advancements" lay immediately 
ahead (Day, 1992, pp. 253, 255): 

Briscoe was soon assigned to other heavy responsibilities. Specifically 
these were under the title of Special Assistant to the President. This role was 
announced by President Wells in May 1939. Then on 1 June 1940 he became 
the first incumbent of the newly created office of Dean of the Faculties. One 
year later, shortly before he was succeeded by a new chairman of chemistry, 
his title was changed to Vice President, Dean of the Faculties. 

122 History of Science: Day Vol. 106 (1997) 

Briscoe was keenly aware of the concurrent developments in the Amer- 
ican Chemical Society on the professional training of chemists. Already a pro- 
gram of the ACS was underway for the certification of institutions for such 
training. Naturally he wanted his department to qualify for certification as 
soon as possible. In essence he worked understandingly with the university 
administration to transform the department. 

Aware of weaknesses in his Department, he moved promptly and wisely. For 
example, on August 26, 1938, Briscoe informed Dean Stout (College of Arts and 
Sciences) that William Degnan, a recent graduate at Yale, had accepted an instruc- 
torship at Indiana University effective immediately. Degnan was the first "out- 
sider" appointed full-time to the chemistry faculty in the 20th Century! 

An illuminating example of Briscoe's administrative genius is found in his 
twenty-page double-spaced report to President Wells on the Department of Chem- 
istry (Briscoe, 1938). The one-paragraph transmittal letter read: 

I enclose herewith a statement concerning the present status and needs of the 
Department of Chemistry. In this I have attempted to outline my evaluation 
of the present department in all respects and to state conservatively my esti- 
mate of its immediate needs. It is my opinion that the department should 
take immediate steps to place itself on a par with some of our neighbors 
such as Northwestern, Nebraska, and Iowa and plan for a future when we may 
rival even our superior neighbors such as Illinois and Wisconsin. I should be 
glad to discuss this report with you at your convenience. 

The report had eight sections. The titles and brief commentaries follow: 

MENT Considering the question of "the organization of chemistry at Indiana 
University as a school instead of a department of the College of Arts and Sci- 
ences," he wrote that: "It is my opinion that the work of the department is too 
intimately connected with the college and with the professional schools to allow 
organization upon an entirely independent basis." After further characteristi- 
cally thoughtful commentary, he stated: 

It is my opinion that the department should be administered through a 
chairman appointed by the President and Board of Trustees for a definite peri- 
od of time with the understanding that reappointment is possible and proba- 
ble in the event of satisfactory service. The limitation placed upon the term of 
office should make change easy when it becomes desirable. The chairman 
should consider his entire staff as a committee for the consideration of the fun- 
damental policies of the department with respect to curriculum, general depart- 
mental business, and student affairs. A good chairman who has sound ideas 
concerning such policies should be able to carry his staff along with him. If 
he cannot, he should be able to subject his own ideas to severe criticism and, 
perhaps, to alter them until they are more nearly in agreement with those of 
the committee as a whole. The chairman should assume the responsibility of 
carrying out the policies and regulations passed on to the department from the 
administration of the university and more particularly from the administra- 
tion of the college. Only in matters which are left to the consideration of the 
department or in dealing with problems which arise in the department, [sic] 
should the department as a whole assume active participation. 

Vol. 1 06 ( 1 997) Indiana Academy of Science 1 23 

The new (less than 90 days) Chairman had focused on his new responsibilities. 
This section was followed by his reflections on the Department's activities 
over the past several decades. 

The first Chairman concluded that "the department has four functions in con- 
nection with its position in the College of Arts and Sciences." He pointed out 
that first: "It must act as a service department for certain professional schools — 
training in chemistry on both the lower and the higher level of the curriculum." 
Second, he concluded that the Department "should offer to all the students of 
the university general courses in chemistry which will fit into the liberal edu- 
cation program of the College of Arts and Sciences." He emphasized that "the 
department should offer a special course to those students." Third, the Depart- 
ment should "provide professional training for the students who will become 
chemists in the industries." Finally, he noted that: "The department must pro- 
vide for original work in the field of chemistry. Provision must be made for a 
modern and adequate program of graduate instruction, for the proper control and 
guidance of the research work done by students in the graduate school, and for 
the scholarly studies, investigative work, and writing of its staff. This function 
should be second to no other function in the department." 

was used to point out the deficiencies in the chemistry curriculum at that time. 
Briscoe summarized this section by saying: "our curriculum stresses the practi- 
cal aspects in chemistry and makes little attempt to provide the necessary require- 
ment for original and scholarly work." Then, he listed some very important 
changes that would have to be made at the undergraduate and master's levels. 
In beginning his analysis of the needs at the upper graduate level, Briscoe wrote 
that the upper graduate level "should be one that is designed for the few .... I 
would not go so far as to say that we should abandon the A.M. degree. We should, 
however, place emphasis upon a curriculum designed for the Ph.D." Briscoe 
wanted these basic but major changes made as quickly as possible. 

Section IV. THE STAFF OF THE DEPARTMENT. The new Chairman wrote 
in effect that in all changes "we must have in mind primarily the graduate work 
in the department and the productiveness in research by members of the staff.*' 
The relatively long discussion devoted to enlarging the chemistry faculty is 
impressive. In the areas of organic chemistry and physical chemistry, Briscoe 
listed 14 persons whom he considered to be "representative of the kind of indi- 
viduals whom I think Indiana University should seek." He expressed the opin- 
ion that "some of them, at least, might find an offer from Indiana University 
attractive." Those named included R.L. Shriner and Henry Gilman in organic 
chemistry and Henry Eyring and Farrington Daniels in physical chemistry. In 
the search for a Chairman to succeed Briscoe so that he could devote all of his 
time to central administration, the University was indeed fortunate that Shriner 
accepted the offer to join the faculty as Professor of Chemistry and Chairman of 
the Department. 

124 History of Science: Day Vol. 106 (1997) 

In this section, Briscoe also referred to the need for additional new faculty 
members at lower academic ranks. Included was a person to teach and do research 
in biological chemistry. He pointed out that this individual would strengthen both 
biology and chemistry. (The appointment also added strength to the move of 
first-year dentistry to Bloomington in 1940-41.) 

Once more, Briscoe emphasized that the persons selected should be "the 
graduates of institutions other than Indiana University." Concerning their cre- 
dentials and promise, he emphasized that: "They should be selected also upon 
their records and promise along the line of research." 

Section V. SPACE. His analysis and recommendations on space requirements 
were based on expectations for the future. Briscoe wrote: "With the growth of 
the department and the expansion of its program more space should be made 
available in the east wing now occupied by the Department of English." Aware 
of the dire needs for improvements in space as well as in general moderniza- 
tion in physical chemistry, he stated that: "The space on the third floor of this 
wing should be converted into a modern laboratory for physical chemistry." 

Section VI. EQUIPMENT. Because he had high expectations for the Depart- 
ment, Briscoe wrote: "At present the equipment is entirely inadequate for under- 
graduate instruction, and there is practically nothing in the department in the 
way of first class modern equipment for research. . . . The appropriations of the 
department in the past has done little more than replace the chemicals, glass- 
ware, and supplies consumed during the previous year." His report on the dire 
needs in the Department reflected the thoroughness of his analysis. 

report, various other matters were discussed. These matters included: "Funds 
from outside the university in support of research" (with elaborations on the 
need); Service to the People of Indiana; Research Directed at the Solution of 
Indiana Problems; and the value of adding "to the department a man trained in 
the field of nuclear chemistry." Briscoe felt that a nuclear chemist would be a 
valuable addition to the Department in view of the research program in physics 
and the possibility of tying these two important fields together. 

Section VIII. LABORATORY FEES. His discussion of the problems in this 
area was concluded with the words: "Increased laboratory fees are recommended 
only in the event that the university is unable to provide for laboratory needs 
from other sources of income." 

(1938) AND PLANS FOR THE PRESENT YEAR. The final section of the Novem- 
ber 1938 report was a listing of different internal actions that would advance the 

1. Representatives of leading companies should be invited to interview 
graduating students for employment. 

2. Tuesday mornings were set aside by the Chairman to talk with stu- 
dents individually in his office to offer "guidance in the formation of 
plans for the future." 

Vol. 106 (1997) Indiana Academy of Science 125 

3. Because a "survey of the curriculum has not been made for many years," 
the Department was considering revisions "along the lines mentioned 
previously in this report." 

4. Attention was being given to ways to "attract some conferences and 
meetings of importance to our campus." 

5. A departmental committee was considering how the present standards 
of graduate work might be changed "to elevate these standards to a high- 
er level of accomplishment." 

Even before Dr. Briscoe succeeded Dr. Lyons, a central tenet of his think- 
ing was that the leadership of the Department had to come from a highly qual- 
ified person with a good national standing in chemistry and with some promise 
of administrative capability. As Briscoe prepared to relinquish this post, he noted 
that his successor's principal education in chemistry and related areas should 
have been obtained elsewhere (Day, 1992, p. 281): 

Characteristically Briscoe moved quietly in searching for his successor. 
He sought and listened to the views of faculty members in the department and 
trusted leaders in chemistry elsewhere. This culminated in the selection and 
attraction of Dr. Ralph L. Shriner, the right person for the times and the needs 
of the department. He was a productive, resourceful, and widely recognized 
leader in both the academic and industrial world of chemistry. 

The public announcement of the selection was made by President Wells 
in the IDS [campus newspaper] of 15 July 1941. . . . The final decision was 
made by Briscoe and Wells in May. The board approved the appointment on 
2 June and the formal notification [to Shriner] was made by Wells a week later. 

In retrospect, one might say this about Briscoe's three-year chairmanship 
(Day, 1992, p. 279): 

He led the university in starting the department toward an ever expand- 
ing level of productivity in chemical education and research. Its stature and 
respect in professional chemistry acquired increasing significance. Within less 
than four decades it became a truly major department in this country. In this 
development all areas of the university shared equally in deriving the bene- 
fits from his wisdom, devotion, and credibility. 

Briscoe's nature and the basis for his overall effectiveness were aptly expressed 
in the words of the trustworthy administrative secretary of his last sixteen years, 
Lucile B. Languell (Day, 1992, p. 280): 

He was not only a great administrator and teacher but he was also a true 
friend to all who sought his valuable advice. He was completely unselfish, 
always thinking of the interests of others, and it was his kindly ways that 
brought so many people to him for help with their problems. Among these 
people were not only faculty members but staff employees as well. 


After Dean Briscoe's principal responsibilities in the Department of 
Chemistry had been transferred to others (the end of the summer of 1941). he 

126 History of Science: Day Vol. 106 (1997) 

discreetly remained aloof from departmental administrative matters, but he 
retained his academic office-laboratory in the chemistry building for several 
years. With pleasing frequency, he came to his office in the chemistry building 
from his main office in Bryan Hall (Administration), walked about in the 
building, and conversed with others. Obviously, he enjoyed being with chemists. 
He also made notable use of the chemistry library, which was directly below his 
office-laboratory. Briscoe's stroke in 1945, the consequent partial health impair- 
ment, and the increasing space needs of the Department ultimately required that 
he surrender his office-laboratory for other uses. 

Between approximately 1941 and 1960, Day (1987, p. 22) noted that: 

Many academic and administrative developments occurred which bore 
the deep imprint of Dean Briscoe's counsel and had his creative participation. 
Several had been included in some degree in the deliberations of the Self-Sur- 
vey Committee. Those issues in which he was notably involved included the 
Junior (University) Division, U.S. War Manpower Commission, School of 
Health, Physical Education, and Recreation (HPER), School of Optometry, 
School of Music, Graduate School, orientation of foreign students for study 
in America, Special Administrative Committee of the Presidency, and Acad- 
emic Freedom. 

In addition to these creative activities, additional responsibilities existed that 
were ongoing and equally demanding — budget preparation and control as well 
as meetings with the President of the University and school deans and depart- 
mental chairpersons in the College of Arts and Sciences. 

Budgets. In the new administrative framework, budget preparation and the 
control of approved budgets became the responsibility of the Dean of the Fac- 
ulties and Vice President. Every academic budget was under Dr. Briscoe's gen- 
eral control, but he exercised his authority so gently and constructively that 
departmental chairpersons and school deans could only conclude that his judg- 
ment was always fair and appropriate. For example, in 1983, then Professor 
Emeritus Harry Sauvain of the School of Business wrote to the author on his 
experiences as Acting Dean of the School of Business (Day, 1987, p. 23): "When 
I went to him with a proposal or a problem he would listen patiently while I told 
him all about it ... I always went away feeling that I had had a fair hearing and 
that the decision was a just one." 

Junior Division. As early as the mid- 1920s, Briscoe believed that thought- 
ful student guidance was basic to education. Typical of his position was a 14- 
page memorandum which he sent to President Wells (written on October 3, 1941). 
The memorandum was "a proposal for the organization and administration of a 
lower division [in the University] consisting of the first year. It is based very 
largely upon the organization which I saw in operation at Yale and upon the orga- 
nization at Nebraska." 

Characteristically, the memorandum was prefaced with an "Introductory 
Statement" in which Briscoe emphasized (1941, p. 1) that, for students, "The 
first and sometimes even the second year are years of adjustment and explo- 

Vol. 106(1997) Indiana Academy of Science 127 

ration; the last years are, and should be, years of concentration in one field . . ." 
Central to the entire presentation (Briscoe, 1941, p. 9) for establishment of a 
lower division was the following summation: 

Higher education must . . . have two primary objectives: 

( 1) to broaden the base of a student's general education and 

(2) to provide training in some specialized field. 

The lower division must promote both of these objectives. It must extend his 
[and her] study into fields that he [and she] has not entered in high school or 
carry on studies previously undertaken to new depths. It must also promote 
the second objective sometimes getting him [or her] started on the ground 
floor of a specialized course, but more often in assisting him [or her] to find 
his [or her] field of specialization... 

These objectives were amplified for the President in ten concluding statements 
labeled "The Essential Provisions of a Lower Division." The final provision 
specified that (Briscoe, 1941, p. 13): "Complete responsibility for the work of 
freshmen must be vested in the administration of the lower division, subject to 
general policies that are established by the general faculty of the university." 

Briscoe's 1941 proposal to President Wells was adopted by the faculty 
after the name was changed to Junior Division. Immediately following the fac- 
ulty action, the Board of Trustees gave their approval (Day, 1987, p. 24): 

The new Junior Division became operational almost at once, with Wen- 
dell W. Wright serving as the first dean. The first students enrolled in May 

1942 Over the years its structure and function remained essentially intact, 

but in 1970 the name was changed to University Division. 


Throughout Briscoe's approximately two decades of administrative ser- 
vice to the University, numerous other special contributions were made by the 
Vice-President and Dean of the Faculties. 

War Manpower Commission. In December 1942, a year after this country 
had entered World War II, Paul V. McNutt, then Administrator of the Federal 
Manpower Commission, requested President Wells to release his long-time friend 
Briscoe for a limited time period to serve in Washington to help formulate poli- 
cies on the use of academic institutions in the war effort. This request was prompt- 
ly granted. In Washington, Briscoe represented all the institutions of higher 
learning in this country which were involved with Federal training programs. 
His work required numerous arduous round trips by train to Washington: in the 
interim, Prof. A.L. Kohlmeier served as Acting Dean of the Faculties. The demands 
placed on Briscoe in Washington heightened. By request, his service to the war 
effort was extended when Dean Briscoe was named Director of the War Man- 
power Training Bureau, placing him in the forefront of postwar educational plan- 
ning. Professor Ford P. Hall relieved Dr. Kohlmeier and served as Acting Dean 

128 History of Science: Day Vol. 106 (1997) 

until Briscoe's duties in Washington ended. The precise length of his service in 
Washington is unclear, but Briscoe's entry in Who's Who in America for 1958- 
59 states that he was a consultant to the Commission from 1942 to 1944. 

School of Optometry. In 1945 in response to requests for the establishment 
of a School of Optometry, President Wells asked Vice-President Briscoe to pre- 
pare a report that would serve as the basis for policy discussions. Within the year, 
Briscoe had made a coast-to-coast survey. After considering all the facts, he con- 
cluded that the University had the responsibility to provide such training and 
that the programs in the School of Optometry would not conflict with those in 
Ophthalmology in the School of Medicine. However, owing to some persistent 
objection by the Indiana Medical Association and some ophthalmologists, Briscoe 
deferred action so all the alternatives could be considered. Ultimately, the issue 
was taken to the Indiana General Assembly, and a bill authorizing the School 
of Optometry was passed almost unanimously. Preprofessional work started in 
1951-52. The first faculty member and essentially the creator of the School 
was Henry W Hofstetter, who arrived in 1952. Until its designation as the School 
of Optometry in 1975, the program operated as the Division of Optometry in the 
College of Arts and Sciences. Until his retirement from administrative respon- 
sibilities in 1959, Dean Briscoe was responsible for the budgetary affairs of 
optometry at Indiana University (Day, 1987, p. 26). 

School of Health, Physical Education, and Recreation. Early in 1945, 
Vice President Briscoe reported that the administration was considering con- 
solidating all physical education programs into a single division under the admin- 
istration of an academic dean (Clark, 1977). This change had been recommended 
by the Self-Survey Committee in 1939 and was approved by the general facul- 
ty in 1940. Although some uneasiness and opposition to the consolidation exist- 
ed, the confidence-inspiring nature of Dean Briscoe aided the change. Support 
for a unified division was strengthened by Wendell Wright of the School of Edu- 
cation, who had also been a member of the Self-Survey Committee. The influ- 
ence of Wright, Briscoe, and Wells contributed to the faculty consensus that 
led to the establishment of the new school in September 1945. Willard W. Patty 
became the first Dean (Day, 1987, p. 26). 

School of Music. Dean Briscoe helped in the re-creation of the School of 
Music. The key element was the appointment of Wilfred C. Bain as Dean of 
the School in 1947. Close communication always existed between Briscoe and 
Dean Bain. These two opposites (in some respects) fit together in a productive 
way that was reminiscent of the match between Briscoe and Wells. The net effect 
was succinctly and poignantly expressed by Dean Bain at the time of Dean 
Briscoe's death in 1960 (Day, 1987, p. 27): 

Herman Briscoe was for me a distillate of human goodness with a wealth 
of wisdom. He was a creative thinker not only as a scientist but also as a pat- 
ternmaker for the education of youth. As an academic elder brother he sought 
to administer unselfishly to the needs of each member of his academic fami- 
ly. As a trusted friend he exemplified the nobility of intellect as he dealt with 
problems outside the field of his immediate interest. 

Vol. 106 (1997) Indiana Academy of Science 129 

The Graduate School. The author noted in 1987 that (Day, 1987, p. 28): 

The three major architects in moving toward a redirecting and vitaliza- 
tion of graduate work were the new President Wells, the new Dean of the Fac- 
ulties Briscoe, and the blunt and determined Dean of the Graduate School 
Fernandus Payne. All three were graduates of Indiana University and native 
Hoosiers. With the ascendancy of Wells to the presidency in 1938 they became 
an effective team, each with unique talents, that operated toward common 

Although these three men recognized the need for greater centralization of direc- 
tion in the various graduate programs, several significant barriers to centraliza- 
tion existed which could not be quickly removed (Clark, 1977, p. 353). 

In 1947, Dean Briscoe presented President Wells with a plan in which as 
much of the graduate work at the University as possible would be centrally coor- 
dinated and some degrees combined to minimize duplication. He proposed that 
the professional schools be granted greater freedom in setting their specialized 
standards. Realizing that this proposal would arouse concern and conflict, Briscoe 
tried to include the changes in planning going on as a result of the approaching 
retirement of Dean Payne. Nevertheless, implementation of the plan was delayed 
due to effective opposition. Some coordination was attained, but full imple- 
mentation did not occur in Briscoe's lifetime. 

By 1989, while Thomas Ehrlich was President of the University, extensive 
centralization between all the campuses of Indiana University had been achieved 
through the creation of the University Graduate School. Heading the program 
was George Walker, Professor of Physics and Vice President for Research and 
Dean of the University Graduate School. These changes required more than four 
decades, surely exceeding the original expectations. The changes that occurred, 
along with many other developments, are monuments to these two sons of 
Indiana — Herman Wells and Herman Briscoe. 

Special Administrative Roles. At times, Dean Briscoe and others were asked 
to substitute for an absent President Wells (Wells, 1980, p. 302): 

Owing to Vice President Briscoe's natural closeness to the presidency of the 
university the frequent absence of the president from the university added to 
Briscoe's responsibilities. In October 1947 this became a major responsibil- 
ity. Near that time an urgent request was made upon President Wells to head 
for some time (six months) the Education and Cultural Branch of the United 
States Military Government in Germany. The University Board of Trustees 
granted the request. To make this practicable the Board designated an 
administrative committee consisting of the President of the Board John Hast- 
ings, and Vice-President Briscoe, Wendell Wright, and Joseph Franklin. Dur- 
ing this long interim many decision-demanding issues naturally arose and 
all were handled well by the committee, but without doubt the extra burden 
on Vice-President Briscoe was heavy. 

During Briscoe's years in administration, the research, publications, and pub- 
lic statements of faculty members were occasionally the basis for criticism which 
in effect was a challenge to responsible academic freedom. "As frequent advis- 

130 History of Science: Day Vol. 106 (1997) 

er to President Wells and a builder of a strong faculty, Dean Briscoe was 
always concerned in the maintenance of the university position" (Day, 1987, 
p. 29). One of the notable statements, which reflected Briscoe's feelings, was 
made by President Wells in a letter on May 16, 1955, regarding the work of Alfred 
Kinsey. The relevant portion was quoted by Clark in 1977 (p. 291): "Indiana 
University stands today, as it has for fifteen years, firmly in support of the sci- 
entific project which has been undertaken and is being carried on by one of its 
eminent biological scientists, Dr. Alfred C. Kinsey." 


On June 30, 1959, and in accordance with the retirement policy which he 
helped establish more than twenty years earlier, Dean Briscoe retired from admin- 
istrative responsibilities. Earlier that month the President wrote to Briscoe: 
"the Board of Trustees of Indiana University approved your administrative retire- 
ment on June 30, 1959 from the position of Vice President and Dean of the 
Faculties, with the understanding that you will continue as Professor of Chem- 
istry and will, in addition, serve as Consultant to the President, on a part-time 
basis." The special letter then continued (Day, 1987, p. 30): 

It is the unanimous feeling of the Trustees of the University that the above- 
indicated approval for administrative retirement is given with the greatest 
reluctance. At the same time the Board extends its most sincere congratula- 
tions and appreciation for the splendid record you have made at Indiana 
University which has resulted in such great achievements for the benefit of 
the institution. The Board also is most happy that you will be able to contin- 
ue in service to the University. 

The pending change in Briscoe's status was known well before his actual 
retirement. In August 1958, as Chairman of the Department of Chemistry, the 
author wrote: "we would be delighted if you would be willing to come back into 
the Department of Chemistry following your retirement next year even if it should 
be on a very limited time basis" (Day, 1987, p. 30). President Wells was also 
informed of the Department's interest. In his response, Wells stated that "some 
of us also have had in mind asking Dean Briscoe to direct the University- wide 
effort in program development or to work in some similar general administra- 
tive capacity" (Day, 1987, p. 31). Thus, after July 1, 1959, the retired Dean and 
Vice President moved to a different office — but just as close to the President's 
Office — as Consultant to the President. There, he thoughtfully prepared com- 
mentaries and made suggestions on many administrative matters. As usual, he 
conferred and counseled with innumerable faculty and administrative colleagues. 

Dr. Briscoe's health remained fairly good during the first year of his retire- 
ment. The winter months were spent with Mrs. Briscoe at their home in Flori- 
da. Indeed, Mrs. Briscoe wrote that in the summer of 1960: "He seemed glowing 
with health and good spirits for the first time in several years" (Day, 1987, 
p. 32). This period of good health continued until September 27, 1960, when 

Vol. 1 06 ( 1 997) Indiana Academy of Science 1 3 1 

he suffered a coronary occlusion in his "retirement office" in Bryan Hall in 
Bloomington. He was promptly taken to the IU Medical Center in Indianapo- 
lis. On October 6, 1960, a second attack occurred, resulting in his death two days 

On October 11, 1960, funeral services were conducted in Sarasota, Florida, 
where the Briscoes had owned a home since 1953. Simultaneous memorial ser- 
vices were conducted in Alumni Hall on the Bloomington campus. Participants 
in this service included the local Berkshire Quartet, Rev. W. Douglas Rae, John 
W. Ashton, Frank T. Gucker, Albert L. Kohlmeier, and the University Singers of 
the School of Music. President Wells and the new Dean of the Faculties Ralph 
L. Collins had gone to Florida to be with the Briscoe family and to attend the 
funeral. The Indiana Daily Student reported on that day that: "Not since the death 
in 1955 of Dr. William Lowe Bryan, President Emeritus of the University, have 
similar honors been paid to a member of the university staff (Day, 1987, p. 32). 

The November 1960 issue of The Indiana Alumni Magazine eulogized Dean 
Briscoe in an appropriate double-page tribute. These memorable words were 
spoken by his long-time close friend and colleague President Wells (Anon., 1960, 
p. 7): 

Dr. Briscoe served the University superbly as student, teacher, and admin- 
istrator. Transcending this brilliant record was the influence of his rare spirit 
upon the University. He was a selfless man, never asking for personal recog- 
nition or power; yet because of his great wisdom and quiet strength his advice 
was eagerly sought by his colleagues. He had an unusual quality of personal 
loyalty and an exceptional capacity for friendship. He will ever live in the life 
of the great University he helped to build and in the hearts of his friends. We 
shall not look upon his likes again. 

In the column "Late News Happenings," a statement appeared that "Plans have 
been announced by the University to establish through gifts an endowed chair 
in chemistry to be known as the Herman T Briscoe Memorial Professorship/' 
The planning and the "soft sell" program resulted in an endowment to which 315 
individuals and corporations made contributions. Dr. Dennis Peters presently 
is the Herman T Briscoe Professor of Chemistry. 

This review of the "Superior Role Model" should logically conclude with 
a discussion of the retirement event and the substance of the remarks and actions 
of that memorable evening. Dean Briscoe's public retirement occurred in the 
Alumni Hall on June 5, 1959, at a banquet sponsored by the Indiana Alumni 
Association. Following dinner, two notable events occurred — the poignant, brief 
address given by the retiring Dean and the presentation of Marie Goth's impres- 
sive portrait of Briscoe to the University (Figure 1). 

His long experience as well as his deep-thinking, problem-solving nature 
were evident in every part of his address. Dean Briscoe focused on the basic 
responsibilities of the University. Following a succinct historic introduction, his 
first penetrating remarks were entitled "We Face a Problem." Briscoe (1959, 
p. 8) stated: 


History of Science: Day 

Vol. 106 (1997) 

Figure 1. Beside the Briscoe portrait (left to right): H.T. Briscoe, Verling Votaw, Briscoe's 
former student and President of the Indiana University Alumni Association in 1959, and 
Ralph L. Collins, who succeeded Briscoe as Dean of the Faculties. 

As a University, we face the problem of how we can best perform our func- 
tion, which we have said is to promote the progress of our society. There are 
many ways in which a university may aid the state and its citizens but, in my 
opinion, we can best perform our function through teaching and research, or 
in some areas such as art and music, by creative work. 

Dean Briscoe (1959, p. 8), as part of this focus, emphasized the importance of 
teaching skills and "how to do a job and how to do it well." When elaborating 
on teaching responsibilities and goals, he stated (Briscoe, 1959, p. 8): 

The student should be made to realize his responsibility to his fellow men and 
to society in general. To properly assume this responsibility, the student 
must have some knowledge and appreciation of the world of science and of 
nature and his own place in it. He must have an acquaintance with the deeds, 
thoughts and dreams of other men as revealed in books, art and music. 

The next topic was "Cultivate a Mind to Think." At this point, Briscoe (1959, 
p. 8) emphasized that "the ultimate goal of higher education should be to culti- 
vate in the student a mind that will and can think for itself. . . ." Briscoe (1959, 
p. 8) gave special attention to the responsibilities of the teacher: "The test of 
good teaching is not found in the grades made by superior students, but in what 
the teacher can do and does do for the average and below average students." 

Vol. 106 (1997) Indiana Academy of Science 133 

Those who knew Briscoe well realized that he believed that good teachers encour- 
age and help their students to learn as independently as possible but that they 
also expect the same students to work in cooperation with others. This point was 
brought out in the section on "Higher Education Needs Re-direction." 

Special emphasis was given to research in the section entitled "Research 
Next to Teaching" (Briscoe, 1959, p. 9): 

And now I should like to speak briefly about research as another way in 
which a university serves society. The importance of research is only second 
to that of teaching. Research will make the world of tomorrow different 
from the world of today and it has made the world of today different from 
the world of the Caesars and the Pharaohs. Research is the search for new 
knowledge and once acquired this knowledge may or not prove useful .... 
Every unknown is a challenge, and the unsolved mysteries of the universe, of 
the atom and the molecule, and of plant and animal bodies are just as chal- 
lenging as traveling beyond uncrossed horizons. 

Briscoe (1959, p. 9) emphasized that: 

It is in the field of basic research that colleges and universities can best 
serve, and it is only natural that the teachers who impart knowledge should 
be most concerned with attempts to discover new knowledge. In the social 
sciences, such as economics, political science, and sociology, research throws 
light upon many of the problems that beset us in business, trade and indus- 
try, in government and in our complex social order. Research in these areas 
is, therefore, closely related to our welfare. 

Briscoe (1959, p. 9) also spoke about "creative effort:" 

They do not solve man's problems of shelter and food and physical well being, 
but they are the food of his soul and spirit; they give him vision and hope; they 
provide him with beauty and satisfaction and contentment. 

In his final statement on research, Dean Briscoe (1959, p. 9) said: 

We cannot, therefore, afford to neglect research. It must be supported by those 
who support higher education, because it is in the universities that much of 
the necessary basic research is done. 

In his closing remarks, which were to be his last before the public, he stated 
(Briscoe, 1959, p. 9): 

To grow old in such an atmosphere is to have the glories of autumn and 
the joys of springtime all in one. It is not easy to leave and it will never be for- 


The author expresses gratitude for the privilege of knowing Dr. Briscoe dur- 
ing his first twenty years on the chemistry faculty — which were the last twenty 
years of Dr. Briscoe's life. Throughout that time, Dr. Briscoe was always avail- 

134 History of Science: Day Vol. 106 (1997) 

able for wise and friendly counseling. There are many others, especially Uni- 
versity Chancellor H.B Wells for his trusted evaluations and encouragement, to 
whom the author is also grateful. The author also appreciates the reviewers' com- 
ments on this article. Finally, the author acknowledges the dedicated and thor- 
ough secretarial efforts of Elizabeth M. Greene. They were essential. 


Anonymous. 1960. Dean Briscoe will ever live at Indiana. Indiana Alumni Mag. 23(2): 6-7. 

Briscoe, H.T. 1931. Qualitative chemical analysis: Principles and methods. D. Van Nostrand Co.,New York, 
279 pp. 

. 1935. The structure and properties of matter. D. Van Nostrand Co., New York, 420 pp. 

. 1938. A report to President H.B Wells on the status and future needs of the Department of Chem- 
istry. 20 pp. (The original carbon copy of the report and the transmittal letter are available in the archives 
of the Department of Chemistry, Indiana University.) 

. 1 94 1 . A memorandum to President Wells for the organization and administration of a lower divi- 

sion consisting of the first year. 14 + 2 pp. (Available in the archives of the Department of Chemistry, 
Indiana University.) 

. 1949. General chemistry for colleges, 4th ed. Houghton Mifflin Co., New York, 751 pp. 

. 1959. Teaching is the most important. Indiana Alumni Mag. 21(9): 8-9. (Farewell speech of Her- 

man T. Briscoe as retiring Dean of the Faculties and Vice President.) 
Clark, T.D. 1973. Indiana University Midwestern pioneer: Vol. II. In mid-passage. Indiana Univ. Press, Bloom- 

ington, Indiana, 429 pp. 
. 1977. Indiana University Midwestern pioneer: Vol. III. Years of fulfillment. Indiana Univ. Press, 

Bloomington, Indiana, 678 pp. 
Day, H.G. 1987. Herman T. Briscoe: Indiana University's beloved chemistry teacher and first Dean of the Fac- 
ulties. Unpub. manu., 46 pp. (Copies are available in the archives of the Department of Chemistry and 

the Indiana University Archives, Bloomington, Indiana.) 
. 1992. The Development of Chemistry at Indiana University, 1829-1991. I.U. Printing Serv., 

Bloomington, Indiana, 668 pp. (Copies are available in the office of the Chairman of the Chemistry 

Department and on file in the archives of the Alumni Office). 
Gaugh, H.F 1959. Retiring Dean Briscoe honored by Arts and Science alumni. Indiana Daily Student, June 

6, 1959. 
Wells, H.B 1980. Being lucky: Reminiscences and reflections. Indiana Univ. Press, Bloomington, Indiana, 

493 pp. 

Proceedings of the Indiana Academy of Science 135 

(1997) Volume 106 p. 135-143 







E.M. Ossom and C.U. Ethothi 

Faculty of Agriculture 

University of Science and Technology 

RM.B. 5080 

Port Harcourt, Nigeria 


C.L. Rhykerd 

International Programs in Agriculture 

Purdue University 

West Lafayette, Indiana 47907 

ABSTRACT: An experiment was conducted to evaluate the influence of K 
fertilizer levels and population density on the fresh weight yield and K con- 
tent of the vines and leaves of the fluted pumpkin. The maximum yield of 
leaves and a high K content occurred at a population density of 40,000 plants/ha 
using K 2 at a rate of 100 kg/ha. 

KEYWORDS: Fluted pumpkin, K fertilization, mineral content, plant densi- 
ty, Telfairia occidentalis Hook., yield. 


The fluted pumpkin (Telfairia occidentalis Hook.), which originated in West 
Africa (Irvine, 1969), belongs to the gourd or calabash family (Cucurbitaceae). 
The fluted pumpkin is a large perennial vine grown as a vegetable crop along 
the edges of the closed forest zone in southern Nigeria. The crop is usually prop- 
agated by seeds obtained from mature gourds of the previous harvest. The gourds 
are split open to extract the seeds a few days before planting. The crop is usu- 
ally grown in the rainy season but is more profitably cultivated during the dry 
season, if the plots are irrigated. 

In southern Nigeria, the fluted pumpkin is either inter-cropped with cassa- 
va or yam, or it may be cultivated alone. Most farmers do not maintain any 
specific plant density or level of fertilization; artificial fertilizers are costly, 
and most peasant farmers cannot afford them. The objective of this study was to 
evaluate the influence of K fertilizer levels and plant density on the yield and 
mineral content of the fluted pumpkin. 

136 Soil and Atmospheric Sciences: Ossom, et al. Vol. 106 (1997) 
Table 1. Treatments as a function of fluted pumpkin density and K fertilizer level. 



Fertilizer Level 



50 kg/ha K 2 from 15-15-15 

T 2 


100 kg/ha K 2 from 15-15-15 

T 3 


133 kg/ha K 2 from KC1 

T 4 


266 kg/ha K 2 from KC1 

T 5 


50 kg/ha K 2 from 15-15-15 

T 6 


100 kg/ha K 2 from 15-15-15 

T 7 


133 kg/ha K 2 from KC1 

T 8 


266 kg/ha K 2 from KC1 


The investigation was conducted in the dry season (November 1988 - July 
1989) at the University of Science and Technology's Research and Teaching 
Farm in Port Harcourt (4° 46' N, 7° 01' E) on a Typic Paleudult soil. The aver- 
age rainfall was 2,000 mm per annum (Food and Agricultural Organization, 
1984). The initial fertility of the soil was: pH, 4.50; total nitrogen, 0.06% (deter- 
mined by the semi-Kjeldahl Method); available P, 22.36 ppm (ammonium molyb- 
date method with absorbance and transmittance measured at 660 nm); exchangeable 
K, 41.48 ppm by flame photometry (Allen, et al, 1974). The experiment used 
a split plot in a randomized complete block design in which two planting den- 
sities and four K fertilizer levels were factorially arranged and replicated four 
times. The planting densities were assigned to the main plots, while K fertiliz- 
er levels were in the subplots. Eight treatments (T l to T 8 ) were replicated 4 times 
in a randomized manner so that treatments had the following plant populations 
with a spacing of 1 m by 1 m; T r T 4 , 20,000 plants/ha (2 seeds/stand); T 5 -T 8 , 
40,000 plants/ha (4 seeds/stand). Each subplot measured 8 m by 4 m with an 
interplot distance of 1 m as well as a 1 m width round the experiment. Seeding 
was done on 25 November 1988, after plowing and disk harrowing. Manual 
weeding was done 3 weeks after planting and, subsequently, at 8, 17, and 25 
weeks after planting. 

Fertilizer Application. Fertilizer was applied at 9 weeks after planting. The 
ring (30-cm diameter) method was adopted at the following rates (Table 1): T t 
and T 5 , 50 kg/ha K 2 from 15-15-15 (N-P 2 5 -K 2 0); T 2 and T 6 , 100 kg/ha K 2 
from 15-15-15 (N-P 2 5 -K 2 0); T 3 and T 7 , 133 kg/ha K 2 from muriate of 
potash (KC1); and T 4 and T 8 , 266 kg/ha K 2 from muriate of potash (KC1). N 
from urea and P from single superphosphate were added to T 3 , T 4 , T 7 , and T 8 to 
bring their levels of N and Pup to those of the NPK 15-15-15 used in T 1? T 2 , T 5 , 
and T 6 . Therefore, only K varied, while similar N and P levels were main- 

Mulching and Watering. Dry grass mulch (Okugie and Ossom, 1988) was 
applied at the base of each crop stand at the rate of about 3 t/ha. Watering was 
done twice a week at the rate of 2.65 mm irrigation/plot for the first 2 weeks after 
planting. Because of the increasing severity of the dry season, the rate thereafter 

Vol. 106(1997) Indiana Academy of Science 137 

Table 2. The distribution of rainfall and irrigation during the course of the experiment. 

No. of Days 




With Up To 

Total Water Use 



0.1 mm 


Nov. 1988 





Dec. 1988 





Jan. 1989 





Feb. 1989 





Mar. 1989 





Apr. 1989 





May 1989 





June 1989 





July 1989 





was increased to 3.98 mm irrigation/plot twice a week but was discontinued on 
March 3, when the first heavy rain fell. Typically, farmers do not water their flut- 
ed pumpkin plots in the dry season and, as a result, lose much of their crop to 
drought. In this experiment, minimal irrigation was provided as a way of possi- 
bly improving crop growth, avoiding crop failure, and increasing crop yield. 
Table 2 shows the rainfall and irrigation distribution during the course of the 

Harvesting and Sample Preparation. The first leaves and vines were 
harvested 7 weeks after planting; subsequent harvests were made every 4 weeks 
(Ossom, 1986) and continued until 27 weeks after planting. Individual plant yield 
was not determined. During harvesting, each vine or branch of a vine along with 
its leaves was cut off about 60 cm from the growing tip, and the total fresh weight 
of both vines and leaves was recorded. A 200 g sample/plot was taken from the 
weighed material, wilted overnight in the laboratory, and bagged. The samples 
were dried in a hot-air oven at 80° C for 5 days. Then, the dried samples were 
ground in a micro-hammermill using a 0.025 mm mesh screen. Analyses for N, 
P, and K (Allen, et al., 1974) were conducted at 7, 11, 19, and 27 weeks after 


Fresh Weight Yield of Leaves and Vines. Low fresh weight yields were 
obtained during the early harvests (7, 11, and 15 weeks after planting). Higher 
yields were recorded during later harvests (19, 23, and 27 weeks after planting), 
which coincided with the period of heavy rainfall. Though the effect of rainfall 
on the treatments was not specifically tested, the low yields at 7, 11 and 15 weeks 
after planting were probably due to the adverse effects of insufficient rainfall 
during that period. Conversely, the higher yields observed at 19, 23, and 27 weeks 
after planting can probably be attributed to the higher rainfall experienced dur- 
ing March. Cumulative fresh weight yield showed significant differences 

138 Soil and Atmospheric Sciences: Ossom, etal. Vol. 106 (1997) 

20 - 

19 - 

18 - 

17 - 

16 - 


15 - 


14 - 
13 - 



12 - 


LSD (P = 0.0 1) = 4.89 


1 1 - 
10 - 

9 - 
8 - 

1 1 — - 

_j 1 1 1 





120 160 200 

Fertilizer level (kg/ha) 



Figure 1 . The effect of K fertilizer and plant density on the fresh weight yield (t/ha) of 
the fluted pumpkin. 

(P = 0.05) between treatments. Least significant difference tests for plant pop- 
ulation density, fertilizer levels, and interaction indicated by A, B, and AB, respec- 
tively (Table 3), showed significant yield differences (P = 0.05) between treatments. 
Cumulative fresh weight yield was higher at a population density of 40,000 
plants/ha than at 20,000 plants/ha. 

Effect of Fertilizer Level on Fresh Weigh Yield. Different fertilizer levels 
did not influence cumulative fresh weight yield. However, a significant increase 
(P = 0.05) in fresh weight yield during harvests 11 and 19 weeks after planting 
was noted. The increased yields might have resulted from increased fertilizer 
availability (K fertilizer was applied 9 weeks after planting) and more efficient 
water use. Singh and Ghosh (1984) noted that graded doses of K increased the 
dry matter yield of maize, cowpea, and wheat. Hassan and Ayoub (1978) noted 
that NPK fertilization leads to increased yield in the onion. 

Effect of Fertilizer by Plant Density Interaction on Cumulative Fresh 
Weight Yield. Increased fertilizer levels did not result in significant fresh weight 
yield increases, but the interaction between fertilizer and plant density did result 
in increased fresh weight yield at 40,000 plants/ha (Figure 1). At 133 kg/ha K 2 0, 

Vol. 106 (1997) Indiana Academy of Science 139 

Table 3. The mean fresh weight yield (t/ha) of vines and leaves of fluted pumpkin as 
affected by plant density, K fertilizer, and harvest interval. 

rreatment 1 

Weeks After Planting 









Mean 3 




















T 3 









T 4 










T 5 









T 6 









T 7 









T 8 









A 2 LSD 4 (P = 0.05) 









B 2 LSD^ 

l (P = 0.05) 










'(P = 0.05) 









•See Table 1. 

2 A, B = Main effects; AB = Interactions. 

3 Means followed by the same letters do not differ significantly at P = 0.01, according 
to Duncan's New Range Multiple Test. 

4 Least significant difference test. 

a plant density x fertilizer interaction was noted that resulted in yield differences 
when treatment means were compared using a least significant difference test 
(P = 0.01). 

Effect of Plant Density on Cumulative Fresh Weight Yield. The highest 
fresh weight per harvest was obtained from T 7 . More succulent, harvestable leaves 
and vines were produced at plant densities of 40,000 plants/ha than at densities 
of 20,000 plants/ha. The maximum fresh weight yield of leaves and vines was 
obtained from 15 to 23 weeks after planting. During this period, more soil water 
from rains was available compared to earlier harvests, when little or no rainfall 
occurred (Table 2). Since better fertilizer use is achieved in the presence of soil 
water, the fluted pumpkin probably made more and better use of the fertilizer 
applied earlier as indicated by the production of more vegetative yield. Plant 
population density exerts a prominent influence on the yield of a crop; hence, 
farmers need to increase the number of plants/ha to maximize growth. For 
most crops, yield is expected to increase with increasing density up to a certain 
point after which yields would decline. At higher densities, more leaves and 
longer shoots tend to be produced by the fluted pumpkin in the plant's effort to 
increase its photosynthetic area that might otherwise be reduced by mutual shad- 
ing from nearby plants and plant parts. The favorable effect of increasing pop- 
ulation density on yield agrees with reports from similar investigations using 
pigeon pea (Cajanus indicus; Abrams and Julia, 1974), cowpea (Vigna unguic- 

140 Soil and Atmospheric Sciences: Ossom, et al Vol. 106 (1997) 

Table 4. N concentration (%) in the leaf blades of the fluted pumpkin as affected by plant 
density, K fertilizer, and harvest interval. 

Weeks After Planting 


Treatment 1 





Std. Dev. 








T 2 







T 3 







T 4 







T 5 







T 6 







T 7 







T 8 












LSD 2 (P = 0.05) 







Std. Dev. 







1 See Table 1 . 

2 Least significant difference test. 

Table 5. P concentration (%) in the leaf blades of the fluted pumpkin as affected by plant 
density, K fertilizer, and harvest interval. 

Weeks After Plantin; 



Treatment 1 





Std. Dev. 








T 2 







T 3 







T 4 







T 5 







T 6 







T 7 







T 8 












LSD 2 (P = 0.05) 







Std. Dev. 







1 See Table 1 . 

2 Least significant difference test. 

Vol. 106 (1997) Indiana Academy of Science 141 

Table 6. K concentration (%) in the leaf blades of the fluted pumpkin as affected by plant 
density, K fertilizer, and harvest interval. 

Weeks After Planting 



Treatment 1 





Std. Dev. 








T 2 







T 3 







T 4 







T 5 







T 6 














T 8 














LSD 2 (P = 0.05) 







Std. Dev. 







1 See Table 1 . 

2 Least significant difference test. 

ulate; Adetiloye, 1986), long beans (Vigna sesquipedalis; Choo, 1974), maize 
(Zea mays; Choudhary, 1981), sunflower {Helianthus annus; Ogunremi, 1979), 
and common bean (Phaseolus vulgaris; Leakey, 1972). 

Because of the sparse rainfall and the small amount of irrigation provided 
during the early stages of the experiment, which was followed by heavy rain- 
fall that improved the water balance within the plant and thus encouraged veg- 
etative growth in the later stages of the experiment, the increased plant density 
probably resulted in more soil coverage, thus reducing evaporation and increas- 
ing yield. Gregory (1988) showed that fertilizer use does promote rapid leaf 
growth thereby enabling plants to cover the surface of the soil and bring about 
a reduction in evaporative losses and an increase in water use efficiency. 

The increased fresh weight yield observed at a density of 40,000 plants/ha 
when compared to that at 20,000 plants/ha was in agreement with observations 
(Thompson and Taylor, 1975) on two cauliflower varieties, "Finney's 1 10" and 
"Kangaroo," in which the yield of the former increased considerably with increased 
planting density. The result also agrees with the findings (Farah, 1975) that yield 
and quality of tobacco were improved as plant density increased. In the present 
experiment, yield increases were probably associated with increased plant den- 
sity and greater vegetative growth at these higher densities; maximum plant den- 
sity has not yet been conclusively determined for the fluted pumpkin. 

Mineral Concentration in the Leaves. At each harvest, irrespective of 
the plant population/ha, the relative concentration of N, P, and K in the leaf blades 
was K > N > P (Tables 4, 5, and 6, respectively). At each harvest, the concen- 
tration of N was generally higher in plants sown at 40,000 plants/ha than in those 

142 Soil and Atmospheric Sciences: Ossom, et al. Vol. 106 (1997) 

sown at 20,000 plants/ha. P concentration did not markedly differ between plant 
populations at each harvest. K content did not vary proportionately with an 
increase or decrease in the level of K 2 applied as fertilizer. The observed 
trend for N accumulation in the vines and leaf blades was in agreement with ear- 
lier reports (Hassan and Ayoub, 1978) that noted that increased mineral content 
in onion resulted from increased NPK fertilization. In the present experiment, 
how crop density contributed to the differences in the amount of N in the 
leaves is not clear. However, the results agree with previous investigations 
(Orluchukwu and Ossom, 1988) that showed a significant difference in the con- 
centration of P and K in the leaf blade of the fluted pumpkin grown under dif- 
ferent management practices. At 27 weeks after planting, yellowing of the leaves 
and laboratory tests indicated a low concentration of N as was also observed at 
7 weeks after planting prior to fertilization. 

The mineral content of a plant organ depends, among other factors, on the 
age of the organ and the presence or absence of elements that can either antag- 
onize or promote nutrient reactions, such as chemical fixation and reduction 
(Brady, 1974). N levels of below 0.05% are indicative of the onset of a defi- 
ciency (Purvis and Carolus, 1964). K levels below 0.3% to 0.5% show the onset 
of a deficiency (Purvis and Carolus, 1964). In this study, mineral levels were in 
the deficiency range prior to fertilizer application, but after fertilizer was applied, 
increased mineral concentration was found in the plant tissues. However, con- 
tinued harvest without additional fertilizer application caused mineral levels to 
decline, giving rise to the deficiency symptoms observed towards the end of the 
experiment. The deficiency symptoms could also have resulted from the matu- 
ration of the pumpkin plants. Purvis and Carolus (1964) noted that N deficien- 
cy might occur during crop maturation. 

The moisture holding capacity of the experimental plots was low for three 
reasons: (1) the sandy loam soil had a low water holding capacity; (2) temper- 
atures during the dry season were high; and (3) desiccating winds were com- 
mon. The available soil moisture was not at an optimum level in the early stages 
of the experiment, including the time when the plots were irrigated. 


The results of this experiment indicate that the yield of leaves and vines 
increases when fluted pumpkin is planted at high densities and fertilized. Max- 
imum yields were obtained at 40,000 plants/ha at a K 2 application rate of 100 
kg/ha. If farmers adopt this planting density and level of K fertilization, they 
should get high yields of leaves and vines from this crop. Though it is costly 
for farmers to obtain large numbers of fluted pumpkin seeds for planting, the 
high profits obtained from sales should offset the cost of purchasing the seeds. 
In the future, the relationship between irrigation and yield of fluted pumpkin 
vines planted during the dry season should be studied. 

Vol. 106 (1997) Indiana Academy of Science 143 


Abrams, R. and F.J. Julia. 1974. The effect of planting time, plant population and row spacing on 

yield and other characteristics of pigeon pea. Trop. Abstr. 29(12): 973. 
Adetiloye, P.O. 1986. Effects of mixture, plant population and two intercropping patterns on the 

performance of maize-cowpea association. Nigerian J. Agron. 1(3): 73-77. 
Allen, S.E., H.M. Grimshaw, J. A. Parkinson, and C. Quarmby. 1974. Chemical analysis of eco- 
logical materials. Blackwell Sci. Publ., Oxford, 565 pp. 
Brady, N.C. 1974. The nature and properties of soils (8th ed.). Macmillan Publ. Co., Inc., New 

York, 639 pp. 
Choo, W.K. 1974. The effect of fertilizer level, plant density and trellis height on vegetative growth 

and green pod production of longbeans (Vigna sesquipedalis (L.) Fruw). Trop. Abstr. 29(2): 

Choudhary, A.H. 1981. Effect of maize populations and row spacing on crop yield. Expl. Agr. 17: 

Farah, S.M. 1975. The effect of plant density and fertilization on the yield and quality of flue- 
cured tobacco. J. Agr. Sci. 84: 75-80. 
Gregory, P.J. 1988. Water and crop growth. In: A. Wild (Ed.), Russell's Soil Conditions and 

Plant Growth, pp. 338-377, John Wiley & Sons, New York, 991 pp. 
Food and Agriculture Organization. 1984. Agroclimatological data. Africa 1. 
Hassan, M.S. and A.T. Ayoub. 1978. NPK fertilization on the yield of onion. Expl. Agr. 14: 29- 

Irvine, F.R. 1969. West African crops. Oxford Univ. Press, London, 272 pp. 
Leakey, C.L.A. 1972. The effect of plant population and fertility level on yield and its components 

in two determinate cultivars of Phaseolus vulgaris. J. Agr. Sci. 79(2): 259-267. 
Ogunremi, E.A. 1979. The effects of plant population on sunflower (Helianthus annus L.) seed 

yield in western Nigeria. Ife J. Agr. 11: 51-55. 
Okugie, D.N. and E.M. Ossom. 1988. Effect of mulch on the yield, nutrient concentration and 

weed infestation of the fluted pumpkin, Telfairia occidentalis Hook. Trop. Agr. (Trinidad) 

65(3): 202-204. 
Orluchukwu, J. A. and E.M. Ossom. 1988. Effect of management practice on weed infestation, 

yield and nutrient concentration of the fluted pumpkin, Telfairia occidentalis Hook. Trop. 

Agr. (Trinidad) 65(4): 317-320. 
Ossom, E.M. 1986. Influence of harvest interval on yield, crude protein, N, P and K contents and 

longevity of the fluted pumpkin, Telfairia occidentalis Hook. Trop. Agr. (Trinidad) 63(1): 

Purvis, E.R. and R.L. Carolus. 1964. Nutrient deficiencies in vegetable crops. In: H.B. Sprague 

(Ed.), Hunger Signs in Crops, 3rd Ed., pp. 245-286, McKay, New York, 461 pp. 
Singh, J.D. and A.B. Ghosh. 1984. Effect of graded doses of potassium on dry matter yield and 

potassium uptake by maize, cowpea, and wheat. Ind. J. Agron. 29(2): 246-248. 
Thompson, R. and H. Taylor. 1975. Some effects of population density and row spacing on the 

yield and quantity of two cauliflower cultivars. Hort. Res. 14: 97-101. 

Proceedings of the Indiana Academy of Science 1 45 

(1997) Volume 106 p. 145-157 





James H. Bandoli 

Department of Biology 

University of Southern Indiana 

Evansville, Indiana 47712 

ABSTRACT: The mating system of spottail darters is a form of resource 
defense polygyny in which females deposit eggs on the undersurfaces of ben- 
thic cavities defended by males. I investigated the (1) effect of nest site size 
on nest site defense by males and on brood size, the (2) timing and duration 
of nest site defense by males, the (3) effect of male size on nest site acquisi- 
tion, and (4) female choice of nest sites and spawning partners. Field exper- 
iments with two sizes of artificial nest sites (tiles) conducted over two breeding 
seasons indicated that large tiles were defended more frequently by larger 
males and contained larger broods than small tiles. In both years, male size 
correlated positively with the total number of eggs defended, although in 
one year the relationship was not statistically significant. Some males defend- 
ed nest sites for as long as 60 days, and six of 17 males that spawned in one 
year sequentially defended multiple broods. In laboratory experiments, male 
size was an important factor in the acquisition and defense of nest sites, and 
females chose the larger of two males as a spawning partner when nest site 
size was held constant. 

KEYWORDS: Etheostoma squamiceps,fema\e choice, male size, nest site 
defense, nest site size, Percidae, sexual selection, spottail darter. 


Resource defense polygyny occurs when a subset of the male breeding pop- 
ulation is able to monopolize resources sought by females (Emlen and Oring, 
1977). In such systems, males typically compete for resources, and intrasexual 
selection promotes characteristics that enhance resource acquisition and defense 
(Darwin, 1871). Male reproductive success may be influenced by the quality 
of the defended resource (Searcy, 1979; Alcock, 1987), the quality of the male 
(Cote and Hunte, 1989; Ryan, 1991), or both (Thompson, 1986), depending on 
the nature and extent of female choice. 

Darters (Teleostei: Percidae) have a variety of reproductive modes, includ- 
ing broadcast, clumping, and clustering of gametes (Page, 1983). Page (1985) 
considered the most derived of these reproductive modes to be egg-clustering, 
a form of resource defense polygyny in which males defend cavities where 
females deposit eggs. This reproductive mode is ideal for studies of sexual selec- 
tion because both competition among males and female choice can occur. 

Aspects of sexual selection have been investigated in several egg-clustering 
percids, including the tessellated darter, Etheostoma olmstedi (Constantz, 1979, 

146 Zoology: Bandoli Vol. 106 (1997) 

1985), the johnny darter, E. nigrum (Grant and Colgan, 1983, 1984), the fantail 
darter, E. flabellare (Knapp and Sargent, 1989), and the waccamaw darter, E. 
perlongum (Lindquist, et ai, 1984). Mating strategies of the spottail darter, 
Etheostoma squamiceps, an egg-clustering species found in southern Illinois, 
western Kentucky, and southwestern Indiana (Page, et ai, 1992), have received 
limited attention. Information on the natural history of E. squamiceps comes 
largely from population studies in Illinois (Page, 1974). The sexes are of approx- 
imately equal size during their first year; thereafter, males become 5-15% larg- 
er than females. Females spawn at 1 year, while males generally become sexually 
mature in their second year; the maximum age of both sexes is 3+ years. Dur- 
ing the breeding season, males defend cavities under solid benthic structures, 
usually rocks. Females attach a single layer of eggs to the ceilings of these struc- 
tures. Clutch size (eggs spawned by a female at a specific time and location), 
based on counts of four aquarium spawnings, averaged 60 eggs (range 20-120; 
Page, 1974). Fractional spawning by E. squamiceps females has not been doc- 
umented, but other members of the genus show this trait (Gale and Deutsch, 
1985; Weddle and Burr, 1991), and fractional spawning may be widespread in 
the Etheostomatini (Hubbs, 1985). For example, the mean number of mature ova 
in 18 breeding females was 110 (range 28-357; Page, 1974), which exceeds the 
average clutch size and supports the possibility of fractional spawning. Males 
remain with the eggs until they hatch (5-15 days, depending on water tempera- 
ture); females leave the nest after spawning. 

Etheostoma squamiceps is classified as Endangered in Indiana (Indiana 
Department of Natural Resources, 1993), where its distribution is limited to 
the extreme southwestern corner of the State. The breeding season lasts from 
mid-March through late May (pers. obs.). Bandoli, et ai (1991) found that 
E. squamiceps will readily use artificial nest sites with no significant differ- 
ence in mean brood size (eggs defended by a male at a specific time; may include 
clutches from more than one female) compared to natural broods under rocks. 
In that study, the mean number of eggs per brood was 368, but nests containing 
over 1,000 eggs were not uncommon, ranging from 5% (Bandoli, pers. obs.) to 
15% (Page, 1974) of all broods examined. The average brood/clutch size ratio 
indicates that larger broods may contain 10 or more clutches. 

The results of field and laboratory studies on the reproductive biology of 
E. squamiceps in southwestern Indiana are reported in this paper with empha- 
sis on factors that influence male reproductive success. Questions asked were: 
Do males that defend larger nest sites have larger broods? How long does a male 
defend a nest site? Does male size influence the acquisition and defense of nest 
sites? Can males defend multiple broods? Do females select males as spawn- 
ing partners on the basis of either male size or nest site size? 


Experiments on nest site selection and reproductive timing were conducted 
at Carpentier Creek, a first-order tributary of the Bayou Creek drainage in Van- 

Vol. 106 (1997) Indiana Academy of Science 147 

derburgh County, Indiana. Carpentier Creek drains suburban areas interspersed 
with bottomland hardwood woodlots. Siltation and debris have reduced the num- 
ber of available nest sites, limiting E. squamiceps densities and promoting the 
use of artificial nest sites (Bandoli, et ai, 1991). 

The purpose of the field experiment was to investigate the response of 
E. squamiceps to variation in nest site size. If nest site size is important in deter- 
mining male reproductive success, larger nest sites should be defended more fre- 
quently, attract more females, and contain more and larger broods. If male size 
is important in nest site defense, larger males should defend larger nest sites. 
Moreover, if females choose males on the basis of size, male size should corre- 
late positively with brood size when nest site size is held constant. 

These predictions were tested using two different lengths of half-cylindri- 
cal ceramic field tile (10 cm inside diameter) as artificial nest sites. Large tiles 
ranged from 14.5 to 16.0 cm in length and provided an average of 170 cm 2 of 
undersurface area for egg deposition, an area large enough to accommodate over 
1,400 eggs. Small tiles ranged from 7.0 to 9.5 cm in length and provided an aver- 
age of 90 cm 2 of undersurface area, enough to hold approximately 700 eggs. 
Pairs of tiles (one large and one small marked for individual identification) were 
placed in the stream with roughly 3-m intervals between pairs. To encourage tile 
use, rocks near tiles were removed. Twenty and 24 pairs of tiles were placed in 
the stream in early March of 1990 and 1991, respectively. Tiles were checked 
biweekly (1990) or weekly (1991) from mid-March through late May for 
E. squamiceps nests and guarding males. 

Darters were captured by placing an aquarium net over one end of a tile and 
flushing any occupants into the net. Standard lengths of captured fish were mea- 
sured prior to returning individuals to their respective tiles. Eggs in small nests 
were counted in the field; large nests were photographed and eggs counted direct- 
ly from projected slides. Since E. squamiceps females deposit new eggs 
around rather than over existing eggs, photographs of nests under the same tile 
on consecutive censuses could be compared to ensure that only eggs deposited 
since the last census were counted. 

In 1991, the reproductive activities of individual males were monitored over 
time by marking males with unique combinations of fin clips and subcuta- 
neous injections of permanent ink. These markings allowed the investigator to 
ask if males could defend multiple broods, how long a male would defend a nest 
site, and if males change nest sites during a breeding season. Comparisons of 
nest site defense, brood size, and use of nest sites by females between large and 
small tiles were performed on combined data from both years. Analyses involv- 
ing male size were performed separately for each year because identification 
of individual marked males was done in 1991 only. Statistics were generated 
using ABSTAT (Anderson-Bell Corp.) and S-PLUS (Statistical Sciences, Inc.). 
Means reported in the text are ± one standard error. 

148 Zoology: Bandoli Vol. 106 (1997) 

Table 1 . Artificial nest site selection by Etheostoma squamiceps. The numbers in paren- 
theses are standard errors. 












Mean number of tiles available 





per census 1 





Number of observations 





% of tiles guarded at least once 





during the reproductive season 

Mean % of tiles guarded per census 









% of tiles with one or more broods 44.2 16.4 66.7 25 
during the reproductive season 

Number of broods 23 11 21 5 

Mean brood size 575.9 335.4 597.1 473.0 

(77.8) (34.0) (56.8) (134.9) 

Number of guarding males captured 31 16 25 11 
(with and without broods) 

Mean standard length of guarding 
males (mm) 

Number of females captured 









1 Twenty and 24 pairs of tiles were 

placed in 

the stream in 

1990 and 1991. 

, respectively. 


In both years, the availability of large and small tiles was similar despite 
temporary losses due to flooding and disturbance by raccoons and humans 
(Table 1). 

Are Larger Nest Sites Defended More Frequently? Males guarded large 
tiles more frequently than small tiles, with over 95% of the large tiles defend- 
ed at least once during the breeding season. Across both years, a mean of 
32.5 ± 3.2% of the large tiles were defended on each of 16 census dates, signif- 
icantly more than the 10.9 ± 3.2% defense frequency for small tiles (paired 
r-test: t = 6.17, df= 15, P < 0.001). 

Vol. 106 (1997) Indiana Academy of Science 149 

Are Females Attracted to Larger Nest Sites? More females were captured 
under large tiles than under small tiles in each year (Table 1); combined data 
from both years showed this pattern to be significantly different from the equal 
use of large and small tiles expected by chance (x 2 = 5.9, df= 1, P < 0.025). 

Do Larger Nest Sites Contain Larger Broods? Of the 60 broods found 
during the two breeding seasons, 44 (73.3%) were located under large tiles. Brood 
size was significantly larger for large tile nests (X = 586.0 ± 48.3, range 85-1632) 
than for small tile nests (X = 378.4 ± 48.2, range 134-781; two sample f-test: 
t = 2.43, df= 58, P< 0.02). 

Do Larger Males Defend Larger Nest Sites? Males captured while defend- 
ing tiles ranged from 55-81 mm in standard length; females captured under tiles 
ranged from 34-70 mm in standard length. Age estimates based on standard length 
(Page, 1974) place most of the males in the three year old cohort, whereas females 
ranged from one to three years. Two sample f-tests showed that the mean stan- 
dard length of males captured under large tiles was significantly larger than 
that of males captured under small tiles in both 1990 (t = 3.23, df= 45, P < 0.002) 
and 1991 (t = 2.34, df= 34, P < 0.02; Table 1). 

Do Larger Males Defend More Eggs? In 1990, 19 males were captured 
while guarding eggs under large tiles. For these males, standard length corre- 
lated significantly with the number of eggs defended (Spearman's rank corre- 
lation: r s = 0.67, P < 0.005). A similar but nonsignificant trend was seen in 1991 
(broods combined for males with multiple broods: r s = 0.40, P < 0.18, n = 13). 

Do Males Defend Multiple Broods? Of 35 males marked in 1991, 18 did 
not defend broods. Seventeen of these males were captured only once and may 
have spawned outside the study area. One male was captured on three different 
census dates and never had any eggs in his nest. Eleven males each defended 
one brood in the study area; two broods were defended by each of three males; 
and three males each defended three broods. All multiple broods were defend- 
ed sequentially rather than simultaneously, and three males with multiple broods 
changed tiles between broods. Males with multiple broods defended nests from 
30 to 60 days. Among spawning males, fitness (total number of eggs guarded 
during the breeding season) ranged from 134 (smallest single brood) to 2,580 
(three sequential broods). 

Do Males Change Nest Sites? Seven of 14 males with multiple captures in 
1991 changed tiles at least once during the breeding season, although two of 
these males defended the same tiles for four and five consecutive weeks, respec- 
tively. The other seven males were consistently found under the same tiles, and 
three were among the six males known to have defended multiple consecutive 
broods. No significant difference existed between the mean standard length of 
changers (X = 12 A ± 2.5) and nonchangers (X = 71.3 ± 2.6; two sample f-test: 
t = 0.40, df= 12, P < 0.69), and the mean number of observations per darter was 
similar for each group (4.4 and 3.5 captures/male for changers and nonchang- 
ers, respectively). 











69.2 (2.0) 






46.8 (3.0) 


150 Zoology: Bandoli Vol. 106 (1997) 

Table 2. Standard lengths (mm) of spottail darters during the first half (15 March - 16 
April) and second half (23 April - 28 May) of the 1991 breeding season. The numbers 
in parentheses are standard errors. 

Parameter n Mean Range 

Males first seen defending tiles 
First half of season 
Second half of season 

Males first seen defending eggs 
First half of season 
Second half of season 

Females captured under tiles 
First half of season 
Second half of season 

An unanticipated finding was that males first seen during the first half of the 

1991 breeding season (early March through mid- April) were significantly larg- 
er in standard length than those first seen in the second half (two sample r-test: 
t = 2.87, df= 35, P < 0.007; Table 2). Similarly, the mean standard length of 
males which first spawned during the first half of that breeding season was sig- 
nificantly larger than that of males which first bred during the second half (two 
sample r-test: t = 2.21, df= 17, P < 0.05). Additionally, the mean standard length 
of females caught under tiles during the first half of the breeding season was sig- 
nificantly larger than that of females caught after this period (two sample r-test: 
t = 2.11, df= 21, P< 0.05). 


Three sets of laboratory experiments were conducted during the 1990- 

1992 and 1995 breeding seasons in 38-L aquaria (50 cm by 25 cm; 30 cm 
deep) with natural substrate and tiles for nest sites. Darters were maintained on 
a 13L: 1 ID photocycle and fed frozen brine shrimp augmented with live benth- 
ic invertebrates from local streams. Water temperature varied from 18°-22° C. 

Male E. squamiceps used in laboratory experiments were determined to be 
in breeding condition based on coloration (Page, 1974) and had a mean standard 
length of 72.1 mm (range 60-83 mm). Females had distended abdomens indi- 
cating the presence of mature eggs and a mean standard length of 56.7 mm (range 
35-72 mm). Standard lengths of most males were in the range expected for age 
3+ (> 64 mm; Page, 1974). All females and most males were used in one trial 
only; any male used more than once was always paired with a different male. 

Experiment 1: Is Male Size Important in Nest Site Defense? In each of 
13 trials, two males (one 5-29% larger in standard length than the other) were 
placed in an aquarium containing a single large tile positioned to the right or left 

Vol. 106 (1997) Indiana Academy of Science 151 

of center by a coin toss. Trials lasted 10 days (1990) or 6 days (1995) with a total 
of 29-30 observations per trial. Observations were made at least twice per day 
(weekends) up to a maximum of six per day between 0730 and 2100 h with at 
least 1 h between observations. At each observation, the position of each male 
was scored as under the tile or on the substrate outside the tile (non- swimming 
darters rest on the bottom). The percent of observations each male was observed 
alone under the tile was determined for each trial; these observations were aver- 
aged across all trials, and a paired Mest was used to compare the mean tile defense 
frequencies of large to small males. In addition, the percent size difference between 
males for each trial was compared to the percent of time the larger male spent 
defending the tile using Spearman's rank correlation statistic. 

Experiment 2: Does Nest Site Size Influence Female Choice of Spawn- 
ing Site When Male Size is Held Constant? In each of 16 trials conducted in 
1991 and 1992, two breeding males of similar size (less than 3% difference in 
standard length) were placed in an aquarium with one large tile (15 cm long) and 
one small tile (8 cm long). A gravid female was introduced within 24 hours. A 
coin toss determined whether the large tile was placed to the right or left of 
center, and the small tile was placed on the opposite side. Previous experi- 
ments showed that spawning in the laboratory occurred from 4 h to several days 
following introduction of the female, making constant monitoring impractical. 
Therefore, all trials were conducted by periodically examining all tiles for eggs 
and noting the positions of all darters in each aquarium. Observations were made 
at least twice daily (weekends and days on which experiments began or ended) 
up to a maximum of five per day. Observations were conducted between 0730 
and 2100 h with at least 1 h between successive observations. Trials varied in 
length (mean duration 4. 1 d) and ended 24 h after a female spawned or after 
5 days without a spawn. The number of observations per trial ranged from 8-26 
with a mean of 13.3. Darter positions were scored as under a specific tile or on 
open substrate. The percent of observations during which each tile was defend- 
ed by a male was determined for each trial, and these values were used to cal- 
culate mean tile defense frequencies for large and small tiles across all trials. 
These defense frequencies were compared using a paired r-test. The tile under 
which spawning occurred was used to indicate female choice. A chi-square good- 
ness of fit test was used to compare the observed distribution of spawning 
locations with the equal spawning under large and small tiles expected by chance. 

Experiment 3: Does Male Size Influence Female Choice of Spawning 
Partner When Nest Site Size is Held Constant? In each of 26 trials conduct- 
ed in 1991 and 1992, two males of different sizes (large males averaged 14.8% 
greater in standard length than small males) were placed in an aquarium with 
two large tiles, one on each side of the aquarium. A gravid female was introduced 
within 24 h. The duration of the trials varied as in Experiment 2 with a mean trial 
duration of 5.1 d. Observations were performed as in Experiment 2 with a mean 
of 19.1 observations per trial (range 8-28). The percent of observations indi- 
vidual darters were under tiles was determined for each trial, and these values 


Zoology: Bandoli 

Vol 106 (1997) 


V — 



















20 - 





5 10 15 20 25 

Size Difference Between Males (%) 


Figure 1 . The relationship between male size (standard length) and nest site defense in 
13 laboratory trials. Each trial had 29 or 30 observations of two males in an aquarium 
with a single nest site (tile). The horizontal axis is the difference in size between the two 
males expressed as a percent of the standard length of the smaller male. The vertical axis 
is the percent of the observations in which the larger male was alone under the tile. 

averaged across all trials for large and small males. Mean tile defense frequen- 
cies of large and small males were compared with a paired f-test. When spawn- 
ing occurred between observations, the male spawning partner was determined 
by noting which male defended the tile before and after spawning. A chi- 
square goodness of fit test was used to compare the observed distribution of 
spawning partners with the equal spawning with large and small males expect- 
ed by chance. 


Experiment 1: Is Male Size Important in Nest Site Defense? Tiles were 
defended by males on 77% of the observations. The large male was alone 
under the tile 61.5 ± 8.9% of the time, significantly more than the mean for small 
males (15.0 ± 5.0; paired t = 3.48, df= 12, P < 0.003). Moreover, the greater 
the size difference between the males, the greater the percent of observations the 
larger male spent alone under the tile (r s = 0.78, P < 0.005; Figure 1). 

Experiment 2: Does Nest Site Size Influence Female Choice of Spawn- 
ing Site When Male Size is Held Constant? Females spawned in 9 of the 16 
trials. Eight clutches were deposited under large tiles; in one trial, two clutches 
were deposited by the same female, the first under the small tile (12 eggs) and 
the second 2 h later under the large tile (37 eggs). Overall, female choice of nest 

Vol. 106 (1997) Indiana Academy of Science 153 

site was significantly different than expected by chance (x 2 = 4.9, df-\, 

Large tiles were defended by a male in 68.4 ±6.1% of the observations, sig- 
nificantly more than the small tiles (36.8 ± 5.5%; paired t = 3.77, df= 15, 
P < 0.002). In trials where spawning occurred, this difference was also apparent 
before eggs were deposited. During this period, males defended the large tiles 
for 53.5 ± 10.8% of the observations versus 22.0 ± 7.6% for the small tiles, a 
significant difference (paired t = 5.57, df=8,P< 0.0006). When the expecta- 
tion of spawning site choice was altered from uniform (1:1) to one based on 
the observed defense frequencies prior to spawning (1:2.3 for small and large 
tiles, respectively), female choice of spawning site did not differ from expecta- 
tions based on observed tile defense frequencies (\ 2 = 1.07, df= 1, P < 0.5). 

Experiment 3: Does Male Size Influence Female Choice of Mates When 
Nest Site Size is Held Constant? Females spawned in 13 of 26 trials. Eleven 
spawnings occurred with the larger male, one with the smaller male, and one 
female spawned with both (160 eggs with the smaller male, three with the larg- 
er male). Female choice of spawning partner was significantly different than 
expected by chance Or = 5.8, df= 1, P < 0.025). 

Large males defended tiles during 63.8 ± 4.8% of the observations compared 
with 45.8 ± 5.5% for small males, a significant difference (paired /-test: t = 3.05, 
df= 25, P < 0.006). Prior to spawning, the difference in tile defense frequen- 
cies of large and small males was not significant (66.3 ± 8.8% and 56.0 ± 10.4%, 
respectively; paired t = 1.03, df= 1 1, P < 0.33). However, females chose the larg- 
er male as a spawning partner significantly more than expected even when the 
expectation was modified by the observed before-spawning male defense fre- 
quency from 1:1 to 1:1.2 for small and large males, respectively (x 2 = 5.04, 


The results from the field experiments indicated that nest site selection does 
affect male reproductive success as measured by the number of eggs defended. 
Large tiles contained both more broods and larger broods than did small tiles. In 
both field and laboratory experiments, males preferred large tiles as nest sites 
despite the fact that small tiles were capable of holding more eggs (at least 
700) than the average brood found under large tiles (586 eggs). However, 
E. squamiceps broods can be much larger than 700 eggs, occasionally exceed- 
ing 1000 eggs (Page, 1974; Bandoli, etal, 1991). The value of larger nest sites 
may be that they permit an occasional large brood and associated higher fitness. 

The finding that larger males generally guard larger tiles suggests that 
male size is important in nest site acquisition and defense, a pattern also seen 
in the fantail darter (Seifert, 1963) and the johnny darter (Grant and Colgan, 
1984). This finding was further supported by the laboratory experiments which 
showed that, when two males competed for the same tile, the larger male spent 

154 Zoology: Bandoli Vol. 106 (1997) 

more time defending the tile than did the smaller male, and exclusive tile defense 
by the larger male increased as the size difference between the males increased. 

Field and laboratory observations indicated that females also prefer large 
tiles as spawning sites. However, these observations may be confounded by the 
fact that, before spawning, large tiles are more likely to be defended by males 
than are small tiles. Therefore, females may select the large tiles based on the 
frequency of tile defense by males rather than tile size. 

When nest sites were of equal size, females preferred those defended by the 
larger male. This result was not confounded by unequal nest site defense fre- 
quencies before spawning and, therefore, may represent a real choice. In field 
experiments, male size varied directly with brood size among males defending 
large tiles, although the correlation was significant in one year only. 

Male size has been shown to be an important parameter for female choice 
in a variety of fish species with breeding systems similar to E. squamiceps, includ- 
ing the redlip blenny (Cote and Hunte, 1989), river bullhead (Bisazza and Mar- 
conato, 1988), and mottled sculpin (Downhower and Brown, 1980). The value 
of male size as an object of choice by females has several potential correlates. 
First, larger males may be selected because they are better egg protectors (Cote 
and Hunte, 1989). Etheostoma squamiceps eggs in abandoned nests are quick- 
ly exploited by egg predators or lost to fungus (pers. obs.), and successful hatch- 
ing may depend on the guarding behavior of the breeding male. Many E. squamiceps 
males showed a high degree of nest site fidelity, unlike E. olmstedi males, who 
abandon nests with large broods to seek new nest sites, leaving brood defense 
to smaller subordinate males (Constantz, 1985). Large males may be better at 
defending a nest, and females that choose them would be rewarded with higher 
fitness via increased egg survival, a pattern seen in the mottled sculpin (Down- 
hower and Brown, 1980). Even if size is not a good indicator of the ability of a 
male to protect a brood, females may still gain from selecting large males if doing 
so results in a larger nest site. Large nest sites afford more room for egg depo- 
sition, an important factor when nest site availability is limited 
(Constantz, 1979). Further, large nest sites also contain larger broods. A female 
that adds eggs to a large brood may minimize the chance of losing her eggs to 
predators or filial cannibalism through the dilution effect (Kodric-Brown, 1983). 

Finally, selection of large males may improve offspring quality. This hypoth- 
esis assumes that male size is an honest indicator of genetic quality (Kodric- 
Brown and Brown, 1984) with increased nutritional condition, better predator 
avoidance, and greater resistance to parasites as possible correlates. The first two 
correlates are difficult to quantify and cannot be addressed here. Resistance to 
parasites (Hamilton and Zuk, 1982) can be discussed. Strange (1992) found that 
86% of 44 E. squamiceps examined from a southwestern Indiana stream were 
parasitized by enterogastric acanthocephalan worms (Acanthocephalus dims) 
with the highest infection rate (100%) among the largest (oldest) darters. Addi- 
tionally, the intensity of parasitism increased with age from 3.6 to 22.6 worms 

Vol. 106 (1997) Indiana Academy of Science 155 

per darter. Therefore, size alone appears to be an unlikely indicator of parasite 
resistance, although larger individuals might be better able to tolerate the unavoid- 
able parasite load. 

Male reproductive success in E. squamiceps is also influenced by the dura- 
tion of nest site defense, a pattern seen in the egg-clustering johnny darter 
Etheostoma nigrum (Grant and Colgan, 1983). Some male E. squamiceps defend- 
ed as many as three sequential broods, which required several weeks of nest 
guarding. The territorial mating system of E. squamiceps and E. nigrum 
imposes minimal mating costs on parental males (Gross and Sargent, 1985), 
making prolonged nest site defense advantageous. As new clutches are added to 
the brood, the time to complete hatching increases, and opportunities for addi- 
tional spawnings occur. 

In the field, the mean standard length of Etheostoma squamiceps males 
that bred early in the reproductive season was larger than those that bred later 
that season, a pattern also seen in the waccamaw darter (E. perlongum; Lindquist, 
et al, 1984). Several potential benefits are associated with early reproduction. 
First, males that acquire nest sites early may have longer spawning periods, allow- 
ing them to recruit more females. Second, early spawning females had a larger 
mean standard length than those spawning later and may be able to produce larg- 
er clutches (Page, 1983). Third, early spawning may decrease competition for 
nest sites with bluntnose minnows, a sympatric cavity-nesting species that begins 
spawning midway through the spottail darter breeding season in southwestern 
Indiana (pers. obs.). Finally, water temperature influences egg development rate 
(Page, 1983), and temperatures of 20°-22° C have been found to maximize embryo 
survival in Etheostoma lepidum (Hubbs, et al, 1969). Variable water tempera- 
tures in shallow streams make temperature prediction difficult, and males that 
acquire nest sites early are ready to spawn when stream temperatures become 
optimal. Females may also produce larger clutches at this time, a pattern seen in 
Etheostoma rafinesquei (Weddle and Burr, 1991). 

The results of this study suggest that male reproductive success in the spot- 
tail darter is a function of (1) acquisition and defense of large nest sites, (2) early 
and prolonged nest site defense, and (3) female choice, all of which are influ- 
enced by male size. However, other factors not addressed in this study might 
also be important. These factors include the intensity of male breeding coloration 
(Kodric-Brown, 1983; Morris, et al, 1995), the length and intensity of courtship 
and/or territorial displays (Grant and Colgan, 1984; Knapp and Warner, 1991), 
and the presence of eggs in the nest (Knapp and Sargent, 1989), any of which 
may influence female choice. These factors not withstanding, size appears to be 
an important factor in determining fitness in males and may be a factor in the 
sex-specific differences in age at maturity. Growth curves for E. squamiceps are 
similar for males and females during their first year but separate thereafter as 
males become increasingly larger than females (Page, 1974). Males may be delay- 
ing reproduction in order to maximize growth. 

156 Zoology: Bandoli Vol. 106 (1997) 


Several individuals provided valuable assistance during the field and labo- 
ratory phases of this research. Chief among them were R. Strange, T. Bacon, 
M. Deen, S. Hicks, K. Lutterbach, R. Stefanich, and D. Phipps. W. Wilding pro- 
vided statistical assistance. The University of Southern Indiana provided labo- 
ratory space and computing facilities. Portions of this project were supported by 
University of Southern Indiana Faculty-Student Research Awards. 


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R.A. Cloyd, C.R. Edwards, and L.W. Bledsoe. TRICHOME 


J. Pichtel, A. Govey, and K. Lukscay. REMOVAL OF LEAD 




H.G. Day. HERMAN T. BRISCOE (1893-1960): A SUPERIOR 


E.M. Ossom, C.U. Ethothi, and C.L. Rhykerd. INFLUENCE OF 





Volume 106, No. 1-2 (1997) 









S.E. Brown and G.R. Parker. IMPACT OF WHITE-TAILED 

T.P. Simon, R.N. Jankowski, and C. Morris. PHYSICAL AND 

1980-1995 67 

J.O. Whitaker, Jr., R. McKenzie, M. Rakow, B. Leibacher, and